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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):561—573. 2001. 


A new species of electric ray, Narcine leoparda, from the tropical 
eastern Pacific Ocean (Chondrichthyes: Torpediniformes: Narcinidae) 


Marcelo R. de Carvalho 


Division of Paleontology, American Museum of Natural History, Central Park West at 79th Street, 


New York, NY 10024, U.S.A.; marcelo@amnh.org 


Abstract.—A new species of electric ray, Narcine leoparda, is described 
from 17 specimens collected from four localities off the southern Pacific coast 
of Colombia, in shallow waters up to 35 m in depth. Narcine leoparda is 
distinguished from all other species of Narcine by its unique dorsal color pat- 
tern, composed of numerous small white to creamy-white spots and/or ocelli 
that are also present on dorsal and caudal fins, over a brown to reddish-brown 
background. Other characters that in combination further distinguish this new 
species include broadly rounded tooth bands that are sub-equal in width, tail 
length shorter than or sub-equal to disc width or length, spiracles devoid of 
external papillae, lateral tail fold originating under first dorsal fin base, and 
second dorsal fin usually slightly greater than first in both height and length 
of base. Narcine leoparda is similar in overall aspect and proportional mea- 
surements to N. vermiculatus, also from the tropical eastern Pacific (Gulf of 
California south to Costa Rica), but dorsal coloration easily distinguishes in- 
dividuals of all sizes of both species, which do not co-occur. Narcine leoparda 


is the third valid species of Narcine from the eastern Pacific Ocean. 


Species of the electric ray genus Narcine 
Henle, 1834 are small to medium-sized ba- 
toids that are more diverse in the tropical 
Indo-west Pacific region, where at least 15 
valid species are known to occur (Carvalho 
et al. 2000). In contrast, only two previous- 
ly described species of Narcine are here ac- 
cepted as valid from the eastern Pacific 
Ocean: Narcine entemedor Jordan & 
Starks, 1895 and Narcine vermiculatus 
Breder, 1928. Narcine entemedor has erro- 
neously been considered a junior synonym 
of N. brasiliensis (=N. bancroftii) by many 
recent authors (Bussing & Lopez S. 1994, 
Castro-Aguirre & Pérez 1996), who have 
followed the otherwise excellent revision of 
Bigelow & Schroeder (1953) (cf. Beebe & 
Tee-Van 1941, McEachran 1995). Narcine 
schmitti Hildebrand, 1948, described from 
a single specimen from the Gulf of Cali- 
fornia, has previously been considered to be 
a valid species as well (Bigelow & Schroe- 


der 1953, Castro-Aguirre & Pérez 1996), 
but was synonymized with N. vermiculatus 
by Carvalho (1999a), who revised the ge- 
nus. A third and very distinctive species of 
Narcine has been collected from shallow 
waters off the Pacific coast of southern Co- 
lombia, and is described below as a new 
species. 


Methods 


Measurements on the holotype and 12 
paratypes were taken with electronic cali- 
pers in a straight line, point-to-point to the 
nearest tenth of a millimeter. Large speci- 
mens requiring measurements of 150 mm 
or more were measured with the aid of a 
steel ruler or tape measure, and are ex- 
pressed to the nearest millimeter. All mea- 
surements are presented in Table 1, and are 
expressed as proportions of total length, fol- 
lowing Carvalho (1999a). Measurements of 


562 


electric rays are subject to discrepancy due 
to preservational variations, and should be 
interpreted with caution if specimens ap- 
pear damaged or distorted (Fechhelm & 
McEachran 1984, Carvalho 1999b). Mea- 
surements are as follows: total length (TL, 
from tip of snout to posteriormost tip at 
mid-caudal height; in millimeters, indepen- 
dent variable from which all proportional 
values are derived); disc width (DW, across 
widest aspect of disc, usually close to level 
of third gill openings); disc length (DL, 
from anterior snout region to greatest disc 
length, lateral to pectoral axil); preorbital 
snout length (PBS, from in between anterior 
level of eyes to anterior margin of snout); 
preoral snout length (POS, from top of low- 
er tooth band to anterior margin of snout); 
prenasal snout length (PNS, from in be- 
tween anterior level of nostrils to anterior 
snout margin); snout to greatest disc width 
(SDW, from anterior snout to level of great- 
est disc width, measured over mid-disc); in- 
terorbital distance (IOD, straight distance 
between inner margins of orbits); eye 
length (EL, between anterior and posterior 
margins of eye); interspiracular distance 
(ISD, between inner margins of spiracles); 
spiracle length (SPL, greatest antero-poste- 
rior distance through spiracle); spiracle 
width (SPW, greatest lateral extent of spi- 
racle); mouth width (MW, distance between 
mouth corners, measured between junction 
of upper and lower labial cartilages on each 
side of jaws); upper tooth band width 
(UTB, width of exposed upper tooth band 
in between posterior margin of lips [formed 
by the upper labial cartilages], close to 
mouth opening); lower tooth band width 
(LTB, width of exposed lower tooth band 
at anterior margin of lips [formed by the 
lower labial cartilages], close to mouth 
opening); nasal curtain width (NCW, width 
of nasal curtain at greatest width below nos- 
trils); nasal curtain length (NCL, length of 
nasal curtain from level of anterior margin 
of nostrils to posterior-most point at mid- 
line of nasal curtain); distance between nos- 
trils (DBN, between inner margins of nos- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


trils); distance between first gill openings 
(FGO, between inner margins of first pair 
of gill openings); distance between last gill 
openings (LGO, between inner margins of 
last pair of gill openings); branchial basket 
length (BBL, between first and last gill 
openings); pelvic fin length (PFL, length of 
pelvic fin from insertion to posterior-most 
point, measured ventrally); pelvic fin width 
(PFW, distance between outer-most corners 
of pelvic fins, from tip to tip, measured ven- 
trally); anterior margin of pelvic fin (AMP, 
greatest extent from insertion to outer-most 
corner of pelvic fin); posterior margin of 
pelvic fin (PMP, greatest extent from outer- 
most corner to posterior-most point of pel- 
vic fin); tail width (TW, extent across base 
of tail at greatest width, measured dorsally); 
height of first dorsal fin (HFD, distance 
from greatest height at apex to mid-base of 
first dorsal fin); length of first dorsal fin 
(LFD, greatest length of base of first dorsal 
fin); height of second dorsal fin (HSD, dis- 
tance from greatest height at apex to mid- 
base of second dorsal fin); length of second 
dorsal fin (LSD, greatest length of base of 
second dorsal fin); length of dorsal lobe of 
caudal fin (LDC, distance from origin on 
dorsal caudal peduncle to posterior-most tip 
of caudal fin); length of ventral lobe of cau- 
dal fin (LVC, distance from origin on ven- 
tral caudal peduncle to posterior-most tip of 
caudal fin); height of dorsal lobe of caudal 
fin (HDC, measured vertically from upper- 
most tip of caudal fin apex to base of dorsal 
lobe on tail); height of ventral lobe of cau- 
dal fin (HVC, measured vertically from 
lower-most tip of caudal fin to base of ven- 
tral lobe on tail); height of caudal fin (HC, 
greatest distance between dorsal and caudal 
fin margins, does not equal HDC + HVC); 
distance between dorsal fins (DBD, dis- 
tance between posterior tip of first dorsal 
fin base and anterior tip of second dorsal 
fin base); distance between second dorsal 
and caudal fins (SDC, from posterior tip of 
second dorsal fin to dorsal origin of caudal 
peduncle); snout to cloaca length (SCL, dis- 
tance between anterior snout margin to or- 


VOLUME 114, NUMBER 3 


igin of cloaca); cloaca to caudal fin length 
(CLC, distance from posterior tip of cloaca 
to posterior margin of caudal fin, equals tail 
length); snout to first dorsal fin length 
(SFD, distance from anterior margin of 
snout to origin of first dorsal fin); electric 
organ length (EOL, from anterior margin to 
posterior margin of electric organ, mea- 
sured ventrally); electric organ width 
(EOW, greatest width of electric organ at 
its mid-length, close to level of third gill 
slit, measured ventrally); clasper length 
(CL, from posterior tip of cloaca to distal- 
most tip of clasper). 

Counts were taken from radiographs, and 
are summarized in Table 2. Usage of me- 
ristic characters also follows Carvalho 
(1999a). Counts include: propterygium ra- 
dials (PRO); mesopterygium radials 
(MES); metapterygium radials (MET); total 
pectoral radials (TPR = PRO + MES + 
MET); pelvic radials (PVR); first dorsal fin 
radials (FDR); second dorsal fin radials 
(SDR); dorsal lobe of caudal fin radials 
(DCR); ventral lobe of caudal fin radials 
(VCR, includes radial situated in between 
dorsal and ventral aspects of caudal fin); to- 
tal caudal radials (TCR = DCR + VCR); 
exposed vertical tooth rows on upper tooth 
band (UTR, corresponds to tooth rows vis- 
ible externally on upper jaw when mouth is 
closed); exposed vertical tooth rows on 
lower tooth band (LTR, corresponds to 
tooth rows visible externally on lower jaw 
when mouth is closed); trunk vertebral cen- 
tra (TC, from first whole distinguishable 
centrum in synarcual to anterior margin of 
pelvic girdle, further explained below); pre- 
caudal vertebral centra (PC, centra from an- 
terior margin of pelvic girdle to origin of 
upper lobe of caudal fin); caudal vertebral 
centra (CC, from first centrum in caudal fin 
to last distinguishable centrum); total ver- 
tebral centra (TV = TC + PC + CC); ribs 
(R, elongated pleural ribs articulating with 
paired hemal spines, located posteriorly on 
disc dorsal to pelvic girdle area). The di- 
vision of the vertebral column into trunk 
and precaudal centra uses the pelvic girdle 


563 


as a landmark because it was not always 
possible to discern monospondylous to di- 
plospondylous transitions from radiographs. 
Radial elements of the pectoral and dorsal 
fins that are joined at base were counted as 
two separate elements. Tooth counts were 
taken under stereomicroscope and follow 
the method outlined in Stehmann (1978), 
where rows are counted following a cranial- 
caudal orientation. Only exposed tooth 
rows were counted, i.e., rows visible on 
tooth bands when mouth is closed (dissec- 
tion is necessary to count internal tooth 
rows because of the strong labial cartilages 
immediately lateral to the tooth bands, and 
dissection was not always possible). Tooth 
counts are expressed as fractions (numera- 
tor designates the number of exposed rows 
on the upper jaws, and the denominator in- 
dicates the same on the lower jaws). 

Comparative material of all species of 
Narcine was used for the present study, and 
is listed in Carvalho (1999a). Institutional 
abbreviations follow Leviton et al. (1985). 
The term “‘preadult’’ is used to indicate 
specimens that are not sexually mature (or 
“‘adult’’). Clasper rigidity was used as an 
indicator of sexual maturity for males, be- 
cause gonadal maturity usually closely fol- 
lows clasper calcification. 


Order Torpediniformes Berg, 1940 
Family Narcinidae Gill, 1862 
Genus Narcine Henle, 1834 
Narcine leoparda, new species 
Figs. 1-4; Tables 1—2 


Holotype.—USNM 222200, 277 mm TL 
adult female, south of Buenaventura, Co- 
lombia, 02°56'N, 078°07'W, O0—16.6 m (O— 
5 fathoms), R/V Cacique (LK 69-29, L. 
Knapp), 20. ix. 1969. (Fig. 1). 

Paratypes.—USNM 222198 (7 speci- 
mens, adult and preadult), 200 mm TL fe- 
male (with two pups), 174 mm TL male, 
174 mm TL female, 110 mm TL female 
(cleared and stained), one dissected speci- 
men (no size data available, but probably 
close to 150 mm TL), south of Buenaven- 


564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Dorsal (A) and ventral (B) view of holotype of Narcine leoparda, n. sp. (USNM 222200, 277 mm 
TL adult female, from south of Buenaventura, Colombia, 02°56’N, 078°07'W, O—16.6 m). 


tura, Colombia, 03°14'N, 077°33'W, 5-7 m, mm TL female, 127 mm TL female, 86 mm 
sta. 357, FAO, 25. I. 1969; USNM 222199 TL male, south of Tumaco (Punta Mang- 
(6, all preadult), 168 mm TL female, 163  lares), Colombia, 01°39’N, 079°02'30”W— 
mm TL female, 146 mm TL female, 130 01°37'30’N, 079°02’00’W, 33.3 m (10 fath- 


VOLUME 114, NUMBER 3 


Fig. 2. 


oms), sta. 343 (1:55 p.m.—2:55 p.m.), R/V In- 
derena (LK 70-12, L. Knapp), 27. x. 1970; 
USNM 222498, 272 mm TL adult female 
(same data as holotype); USNM 222500, 
230 mm TL adult male, 01°35'05’N, 
079°03'00"”W—01°37'00"N, 079°04'00"W, 


565 


Dorsal (A) and ventral (B) view of paratype of Narcine leoparda, n. sp. (USNM 235919, 206 mm 
TL adult male, south of Tumaco, Colombia, 01°39'N, 079°02'30”"W—01°37'30"N, 079°02'00"W, 33.3 m). Caudal 
fin is downturned in (B). 


south of Tumaco (Punta Manglares), Co- 
lombia, 9.15 m, sta. 342, R/V Inderena, 
(LK 70-11, L. Knapp), 27. x. 1970; USNM 
235519, 206 mm TL adult male, data as for 
USNM 222199. 

Diagnosis.—A species of Narcine distin- 


566 


guished from all others by its unique dorsal 
coloration composed of numerous small 
white to creamy-white spots and/or ocelli 
on disc and dorsal and caudal fins, over a 
reddish-brown or brown background. Ad- 
ditional characters that in combination di- 
agnose this new species include: tail length 
shorter than disc width or length, upper and 
lower external tooth bands subequal in 
width, spiracles with smooth rims devoid of 
external papillae, second dorsal fin usually 
slightly larger than first in both height and 
length of base, and lateral tail folds origi- 
nating under level of first dorsal fin base. 

Description.—Measurements and counts 
are summarized in Tables 1 and 2, respec- 
tively. 

External morphology: Disc rounded to 
somewhat oval, generally about as wide as 
long. Disc width is somewhat more variable 
than disc length (see standard deviations in 
Table 1). Disc overlaps origin of pelvic fins 
only slightly, and with a very small free 
lobe posteriorly where it contacts sides of 
trunk. Greatest width of disc is just poste- 
rior to its mid-length, but somewhat vari- 
able in specimens. Snout broadly rounded 
anteriorly. Preorbital snout corresponds to 
about one-third in disc length. Electric or- 
gan originating just anterior to level of eyes 
dorsally and at level of nostrils ventrally. 
On ventral side, electric organs extend to 
beyond level of last gill-slit; electric organs 
difficult to discern in dorsal view. Gill-slits 
small, only slightly curved and in more or 
less straight line from first to last gill slit. 
Distance between last gill-slits generally 
less than branchial basket length. Spiracles 
and eyes adjacent, without a distinctive sep- 
aration. Spiracles with somewhat developed 
rims devoid of papillae, circular to oval, 
and usually slightly longer than wide. Eyes 
relatively small, about as long as spiracles. 

Nasal curtain wider than long (Fig. 3C), 
with more or less straight posterior margin, 
but faintly developed central lobe present in 
some specimens. Nostrils small and circu- 
lar; distance between nostrils slightly great- 
er than nasal curtain width at posterior mar- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


gin. Prenasal snout length generally less 
than preorbital snout length. Mouth about 
as wide as distance between nostrils. Upper 
and lower tooth bands more or less equal 
in width and both circular in outline. Teeth 
in 12/8—16/12 exposed rows in specimens 
about 175 mm total length or larger, but a 
146 mm total length female has 11/13 ex- 
posed teeth rows. Teeth relatively small, set 
in quincunx arrangement and with single, 
sharp cusp, even on small specimens (e.g., 
146 mm TL specimen); inner rows with 
very sharp teeth. Crowns wider-than-long at 
bases; tooth bases roughly sub-circular. 
Posterior lobes of pelvics more or less 
straight, with only a small free lobe where 
they contact claspers or lateral aspect of 
trunk (in females). Lateral corners of pel- 
vics blunt, not acute. Pelvics generally ex- 
tend from underneath posterior disc region 
to underneath beginning of first dorsal fin 
base. Claspers of larger males extend pos- 
teriorly to close to posterior aspect of first 
dorsal fin base, but do not project greatly 
beyond pelvic fins. Tail relatively short as 
measured from cloaca, shorter than snout to 
cloaca length, and much shorter than disc 
width and length. Tail sub-circular in cross 
section; its width at base moderately broad 
and somewhat variable. Lateral tail fold ex- 
tending as a thin keel in a straight line lat- 
erally on tail, from level of mid- to poste- 
rior first dorsal fin base to lateral aspect of 
caudal peduncle. Second dorsal fin only 
slightly taller than first dorsal fin, and with 
a slightly longer base. Dorsal fins similar in 
shape, and with a small free lobe posteri- 
orly (Fig. 3B). Caudal fin with angular 
apex, relatively tall and not elongated, and 
with more or less straight posterior margin. 
Length of dorsal and ventral lobes of caudal 
about equal, but dorsal lobe slightly taller 
than ventral lobe. Distance between second 
dorsal and caudal fins generally slightly 
greater than distance between dorsal fins. 
Sensory pores indistinct, few in number, 
and scattered on ventral snout area and 
along margins of electric organs both dor- 
sally (hyomandibular canal) and ventrally 


VOLUME 114, NUMBER 3 567 


Table 1.—Morphometric comparisons for Narcine leoparda, n. sp. n is number of specimens from which 
means and standard deviations (SD) were calculated and include specimens greater than 110 mm TL. See 
Methods section for abbreviations. Holotype: USNM 222200; Paratypes: USNM 222198, USNM 222199, USNM 
222498, USNM 222500, USNM 235519. 


Holotype Paratypes n BG SD 
TL (mm) 277.0 110.0—272.0 13 — — 
DW (%) 48.7 46.6—57.3 13 52.0 3.5 
DL (%) 53.1 50.0-58.2 13 53.4 DU) 
PBS (%) 13.3 12.7-15.7 13 14.6 1.0 
POS (%) 14.7 14.5-17.5 13 15).5) 1.1 
PNS (%) 123 12.0-14.4 13 13.0 0.9 
SDW (%) 31.8 21.1-41.0 13 323} 3.0 
IOD (%) 6.4 6.0-8.5 13 7A 0.8 
EL (%) DS 1.8—3.0 13 2.4 0.4 
ISD (%) 6.0 5.8—8.5 13 71 0.8 
SPL (%) Zee 2.1—3.0 13 2.4 0.3 
SPW (%) 1.6 127 13 2.1 0.3 
MW (%) 5.3 5.3—6.7 13 5.9 0.4 
UTB (%) Dod) 2.3-3.2 13 2.6 0.3 
LTB (%) 2.1 1.9-2.8 13 2.3 0.3 
NCW (%) 5.0 5.1—6.6 8 DoT 0.7 
NCL (%) 1.5 1.0—2.4 11 1.8 0.4 
DBN (%) 5.6 5.5=7.3 13 6.1 0.5 
FGO (%) 12.3 11.5-14.5 13 13).3) 0.8 
LGO (%) 6.9 5.9-9.5 13 Vall 0.9 
BBL (%) 9.1 7.2-11.0 13 9.8 1.0 
PFL (%) 19.9 17.8=23.5 13 20.7 1.9 
PFW (%) 26.7 28.7—36.8 13 33.5 3.0 
AMP (%) 12.7 11.3—14.2 13 12.4 0.8 
PMP (%) 19.0 15.5—23.2 13 ORS 2.6 
TW (%) 15.4 16.6—21.6 13 18.9 1.8 
HFD (%) 7.8 6.8—9.3 13 8.2 0.8 
LFD (%) 6.3 4.5-7.6 13 5.9 0.7 
HSD (%) 8.8 7.5—9.9 13 8.8 0.6 
LSD (%) 6.1 5.5—7.3 13 6.2 0.5 
LDC (%) 11.8 11.8-13.1 13 12.3 0.4 
LVC (%) 11.0 11.6-14.1 13 27 1.0 
HDC (%) 4.7 3.2—5.0 13 4.4 0.5 
HVC (%) 3.1 2.24.7 13 3.7 0.8 
HC (%) QZ 7.5-12.5 13 10.3 1.3 
DBD (%) 2D 2.2-3.4 13 2.6 0.4 
SDC (%) 32) 2.5-3.9 13 3}.3} 0.5 
SCL (%) 53.1 50.0-54.8 13 52.6 1.3 
CLC (%) 43.3 39.7-47.0 13 42.6 2.0 
SED (%) 65.0 65.4—70.1 13 68.0 1.6 
EOL (%) 24.2 22.8—29.2 13 25.0 1 
EOW (%) 10.1 9.8—-12.1 13 10.7 0.7 
CL (%) — 15.7-—17.0 3 16.2 0.7 


(ampullary pores only). Few sensory pores Lateral tail fold pores are very scattered and 
dorsally in more or less straight rows on not numerous. Pores on ventral snout area 
snout region. Pores of lateral canal on lat- in more or less parallel antero-posterior 
eral aspect of tail running dorsal to lateral rows, not extending posteriorly beyond ley- 
tail fold, apparently in a straight line and_ el of nostrils. 

terminating on lateral aspect of caudal base. Coloration: In preservative, dorsal back- 


568 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Meristic features of Narcine leoparda, n. sp. A) USNM 222200 (holotype); B) USNM 222500; C) 
USNM 222498; D) USNM 235519; E) USNM 222199; F) USNM 222199. B-F represent paratypes. Dashes 
represent counts not available in radiographs. See Methods section for abbreviations. 


A B Cc 
TL (mm) ZN 230 Dei; 
PRO 18 17 17 
MES 6 6 6 
MET 11 10 12 
TPR 35 33 35 
PVR 19 17 19 
FDR 8 8 6 
SDR 10 9 7 
DCR 26 17 23 
VCR 2H 2p) 24 
TCR 53 39 47 
UTR 16 15 16 
LTR 9 12 13 
TC 28 24 24 
PC 59 59 59 
CC 24 20 23 
TV 111 103 106 
R 8 8 7 


ground coloration composed of a light, 
dark, reddish- or purplish-brown back- 
ground, with white to creamy-white spots 
or ocelli covering most of disc region pos- 
terior to eyes, and extending posteriorly 
over tail. Dorsal and caudal fins also with 
small white spots on all specimens. Spots 
and ocelli generally not fused into irregu- 
larly shaped blotches, especially in smaller 
specimens where spots are clearly separated 
from each other (Fig. 4). Spots are round to 
slightly oval, varying in definition from 
sharply defined to somewhat fuzzy. Spots 
may be present between eyes, especially in 
smaller specimens, or anterior to eyes over 
dorsal snout region. Smaller specimens 
with spots ranging in size from small (about 
equivalent to diameter of eyes and spira- 
cles) to slightly larger (about twice eye-di- 
ameter). Larger specimens have more var- 
iable coloration, with ocelli closely (holo- 
type; Fig. 1) or loosely packed together 
(paratype USNM 235919; Fig. 2). Ocelli 
composed of central white to creamy-white 
spot, surrounded by ring of darker brown. 
Dorsal coloration mostly restricted to back- 
ground coloration in two large paratypes 


D E F Range 
206 163 168 163-277 
17 — 18 17-18 
5 — 6 5-6 

9 — — 9-12 
31 — — 31-35 
19 20 19 17-20 

8 — 8 6-8 
10 10 10 7-10 
21 — — 17-26 
DS) — — 22-27 
46 — 39-53 
12 13 11 11-16 

9 10 8-13 
26 28 24 24-28 
59 60 59 59-60 
22 24 23 20-24 

107 112 106 103-112 
8 7 8 7-8 


(USNM 222498 [Fig. 3A] and USNM 
222500), but remnants of ocelli are visible, 
and small white spots remain present on 
dorsal and caudal fins. Ventral coloration 
uniform creamy-white, but some larger 
adults may have the posterior disc and pel- 
vic fins outlined in light brown. 

Skeletal features: Examination of skele- 
tonized and cleared and stained specimens 
(both from USNM 222198) reveal that Nar- 
cine leoparda shares with other species of 
Narcine, including the type-species N. bras- 
iliensis (von Olfers, 1831), the following 
features indicative of generic monophyly 
(Carvalho 1999a): fused hypobranchial 
plates with sinuous external margins artic- 
ulating with ceratobranchials 2—4, slender 
posterior projection of the heart-shaped bas- 
ibranchial copula, separation of ceratohyal 
and “‘hypobranchial 1” by a small gap, and 
presence of a separate facio-palatine fora- 
men in the orbit. In general aspects, the 
skeletal anatomy of Narcine leoparda is 
similar to that of N. brasiliensis, with rel- 
atively wide and proximally branched an- 
torbital cartilages that bear numerous digi- 
tiform projections distally. Lateral rostral 


VOLUME 114, NUMBER 3 569 


Fig. 3. A) Dorsal view of paratype of Narcine leoparda, n. sp. (USNM 222498, 272 mm TL adult female, 
from south of Buenaventura, Colombia, 02°56’N, 078°07’W, 0—16.6 m); B) lateral tail region of USNM 222498: 
C) nasoral region of USNM 222498. Note that caudal fin is downturned in (A). 


570 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Paratype of Narcine leoparda, n. sp. (USNM 222198, 174 mm TL male, south of Buenaventura, 
Colombia, 03°14'N, 077°33'W, 5—7 m). 


processes are also present anteriorly adja- 
cent to the rostral fenestra of neurocranium, 
providing further endoskeletal support to 
anterior margin of snout. Dorso-lateral pro- 
jections of the nasal capsules (‘nasal cap- 
sule horns”? of Carvalho 1999a) are also 
present. One pair of triangular labial carti- 
lages on each side of jaw symphysis give 
support to “‘lips’’ surrounding external 
tooth bands. Propterygium is subdivided 
into four segments; mesopterygium is 
small, slender and antero-laterally directed, 
and the metapterygium is subdivided and 
inconspicuous. Pelvic girdle with two pre- 
pelvic processes. with expanded, flattened 
tips, and posteriorly curved iliac processes. 

Etymology.—leoparda, modified from 
leopardus (Latin for leopard), in reference 
to its distinctive dorsal color pattern; inter- 
preted as a feminine noun. 

Geographical distribution.—Narcine 
leoparda is known from four localities on 
the continental shelf of Colombia, south of 
Buenaventura and Tumaco (off Punta 
Manglares), in near-shore waters ranging 
from 5 to 33 m in depth. 


Discussion 


Comparisons with congeners.—Only two 
other valid species of Narcine occur in the 
eastern Pacific Ocean: N. entemedor and N. 
vermiculatus. Among these, N. leoparda is 
most similar to N. vermiculatus Breder, 
1928, which occurs farther north along the 
continental shelf of Central America (reach- 
ing as far south as Costa Rica; McEachran 
1995, Carvalho 1999a). Both species are 
clearly distinct in dorsal color pattern. Nar- 
cine leoparda possesses white to creamy- 
white spots that may be substituted by ocel- 
li in larger specimens, with spots not fused 
into larger blotches or vermiculations, as in 
N. vermiculatus, and generally smaller than 
the markings present in the latter species. 
The spots in Narcine leoparda are present 
even on newly born individuals or pups ex- 
tracted from uteri (USNM 222198). Newly 
born specimens of N. vermiculatus, how- 
ever, clearly display laterally elongated 
blotches or stripes and vermiculations that 
are characteristic of this species (e.g., Bee- 
be & Tee-Van 1941, fig. 1, pl. 1; Carvalho 


VOLUME 114, NUMBER 3 


1999a). In N. leoparda, the spots may be 
lost in larger specimens, as USNM 222500 
and USNM 222498 have more uniform dor- 
sal coloration without clearly defined spots 
over disc. However, small white or creamy- 
white spots are still present over dorsal and 
caudal fins, and closer inspection reveals 
that both specimens previously had dorsal 
spots over the disc as well. Narcine leopar- 
da also differs from N. vermiculatus in hav- 
ing lateral tail folds that begin underneath 
first dorsal fin base (either at its mid-length 
or slightly farther posteriorly), and not in 
between the dorsal fins. Other differences 
that further separate both species are disc 
proportions. Disc is generally more rounded 
in Narcine leoparda, with a greater mean 
disc length (53.4% of TL), compared to N. 
vermiculatus (48.4% of TL), which is also 
reflected in relative origin of first dorsal fin 
(originating farther posteriorly in Narcine 
leoparda {mean distance between snout and 
first dorsal origin is 68.0% of TL; compared 
to 64.1% of TL for N. vermiculatus]). Both 
species also have disjunct, allopatric distri- 
butions, as N. vermiculatus has not been 
collected south of Costa Rica. 

Narcine leoparda is easily distinguished 
from N. entemedor, with which it co-oc- 
curs, in presenting spiracles devoid of pa- 
pillae, in morphometric proportions, size of 
adults and coloration (data for N. entemedor 
from Carvalho 1999a). Narcine entemedor 
generally has an olivaceous brown back- 
ground color, usually with dark blotches on 
disc of larger specimens, and symmetrically 
arranged ocelli in smaller specimens. These 
ocelli, however, are very different from 
those present in the holotype of N. leoparda 
in both color and arrangement (ocelli in N. 
entemedor have dark center and are fewer 
in number, as only four or five are usually 
present on disc). N. entemedor is a much 
larger species (up to 750 mm TL) compared 
to N. leoparda (largest specimen is 356 mm 
TL). Males of N. entemedor become sexu- 
ally mature only between 340 and 370 mm 
TL (cf. to 180 mm TL for N. leoparda, see 
below). Numerous specimens of N. ente- 


571 


medor, and of N. vermiculatus, have been 
thoroughly examined from throughout their 
respective ranges, and are further described 
in Carvalho (1999a). 

Narcine leoparda is easily distinguished 
from most other species of Narcine in mor- 
phometric proportions (20 species of Nar- 
cine are recognized as valid in Carvalho 
1999a). In N. leoparda, tail length (as mea- 
sured from cloaca) is subequal to, and gen- 
erally less than, disc width or length, but 
tail length is much greater than the disc in 
the five Australian species of Narcine (N. 
tasmaniensis, N. westraliensis and three un- 
described species) and in N. rierai fom the 
western Indian Ocean. Both upper and low- 
er tooth bands are broadly rounded and of 


subequal width in N. leoparda, contrasting 


to two southeast Asian species (N. brevi- 
labiata and an undescribed species) in 
which the tooth bands are roughly triangu- 
lar in shape, and in which the upper tooth 
band is much wider than the lower. In N. 
leoparda, the second dorsal fin is subequal 
to, or slightly greater than, the first dorsal 
fin in height and length of base, separating 
it from two undescribed western and north- 
ern Indian Oceans forms, in which the first 
dorsal fin is greater than the second. Both 
western Atlantic species (NV. bancroftii and 
N. brasiliensis) also have spiracles with nu- 
merous papillae, as in N. entemedor, and 
are therefore easily distinguished from UN. 
leoparda. The remaining five species of 
Narcine (N. timlei, N. maculata, N. lingula 
and two undescribed species from the 
northern Indian Ocean and Indonesia) are 
distinct from N. leoparda in coloration, as 
they have either elaborate dorsal patterns 
composed of brown spots and/or blotches 
over a lighter background, or a uniform col- 
oration (N. timle?). 

Biological notes.—One partially dissect- 
ed female of 277 mm TL (holotype, USNM 
222200) had a large egg mass in the right 
uterus, while a female of 200 mm TL 
(USNM 222198) contained two small late- 
term pups in the left uterus. Both pups are 
female, just slightly over 50 mm TL, and 


572 


the smallest one still has remnants of a 
yolk-stalk. No external gill-filaments or ex- 
ternal teeth are present, but few internal 
teeth with small cusps can be observed. 
Both specimens were probably very close 
to birth. One small male of 127 mm TL still 
has a fading yolk-scar and claspers that do 
not project beyond the distal tips of pelvic 
fins. Males probably mature at around 180 
mm TL, as a 174 mm TL male has claspers 
almost completely firm. There are 10 spiral 
valve turns in the intestine of at least one 
specimen examined (USNM 222198, 
cleared and stained). Small, unidentified 
ctenoid scales were present in the mouths 
of two specimens (USNM 222500 and 
USNM 222199, 168 mm TL), and therefore 
N. leoparda feeds also on fishes as do other 
species of the genus. 


Key to the Eastern Pacific species of 
Narcine 


la. Spiracles with papillae present on outer 
GAT Perce eu tees ee acne ee ee N. entemedor 
1b. Spiracles devoid of any papillae 
2a. Dorsal coloration composed of horizon- 
tally elongated, white or creamy-white 
stripes and blotches ..... N. vermiculatus 
2b. Dorsal coloration with small white or 
creamy-white spots and/or ocelli, or rel- 
atively uniform in larger specimens with 
perhaps few spots over dorsal and caudal 
fins, but never with horizontally elongat- 
ed stripes and blotches ...... N. leoparda 


Acknowledgements 


The staff of the USNM was extremely 
helpful in numerous ways during research 
related to this paper, in particular Jerry Fin- 
an, Lisa Palmer, Leslie Knapp, Susan Jew- 
ett and Lynne Parenti, and are thanked for 
their hospitality. David Catania, John Fong, 
Tomio Iwamoto and Bill Eschmeyer (CAS), 
and John Moore (formerly at YPM) also 
provided assistance during visits. Portions 
of this paper are contained in my PhD dis- 
sertation, and I thank my doctoral commit- 
tee (especially John D. McEachran) and the 
Department of Ichthyology of the AMNH 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


for support during the tenure of my studies 
there. Financial support was provided by a 
fellowship from the Conselho Nacional de 
Desenvolvimento Cientifico e Tecnolégico 
(CNPq) of the Brazilian Federal Govern- 
ment, a graduate student fellowship from 
the Office of Grants and Fellowships of the 
AMNH, the USNM (short-term visitor), the 
Department of Ichthyology of the AMNH 
and The City University of New York. All 
institutions are gratefully acknowledged. 
This paper was completed during a post- 
doctoral appointment in the Division of Pa- 
leontology (AMNH), and I thank John 
Maisey and Herbert R. and Evelyn Axelrod 
for the opportunity. 


Literature Cited 


Beebe, W., & J. Tee-Van. 1941. Fishes from the trop- 
ical Eastern Pacific (From Cedros Island, lower 
California, south to the Galapagos Islands and 
norther Peri.) Part 3, Rays, Mantas and Chi- 
maeras.—Zoologica (26):245—270. 

Bigelow, H. B., & W. C. Schroeder. 1953. The Fishes 
of the Western North Atlantic, Part Il. Sawfish- 
es, Skates, Rays and Chimaeroids—Memoir of 
the Sears Foundation for Marine Research, 2: 
xv + 1-588 pp. 

Bussing, W. A., & M. I. Lopez S. 1994. Demersal and 
pelagic inshore fishes of the Pacific coast of 
lower Central America. Pp. 32—33 in Special 
publication of the Revista de Biologia Tropical. 
Escuela de Biologia y CIMAR, Universidad de 
Costa Rica. 

Carvalho, M. R. de. 1999a. A systematic revision of 
the electric ray genus Narcine Henle, 1834 
(Chondrichthyes: Torpediniformes: Narcinidae), 
and the higher-level relationships of the orders 
of elasmobranch fishes (Chondrichthyes). Un- 
published Ph.D. dissertation, The City Univer- 
sity of New York, 735 pp. 

. 1999b. A synopsis of the deep-sea genus Ben- 

thobatis Alcock, 1898, with a redescription of 

the type-species Benthobatis moresbyi (Chon- 
drichthyes: Torpediniformes: Narcinidae). Pp. 

231-255 in B. Séret and J.-Y. Sire, eds., Pro- 

ceedings of the 5th Indo-Pacific Fishes Confer- 

ence, Nouméa. Paris. 

, Compagno, L. J. V., & P. R. Last. 2000. Fam- 

ily Narcinidae. Pp. 1432—1442 in K. Carpenter 

and V. Niem, eds., FAO Species Identification 

Guide for Fisheries Purposes. The Living Ma- 

rine Resources of the Western Central Pacific. 

FAO, Rome. 


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Castro-Aguirre, J. L., & H. Espinosa Pérez. 1996. Lis- 
tados faunisticos de México. VII. Catalogo sis- 
tematico de las rayas y expecies afines de Méx- 
ico. (Chondrichthyes: Elasmobranchii: Rajifor- 
mes: Batoideiomorpha).—pp. 7—75. 

Fechhelm, J., & J. D. McEachran, 1984. A revision of 
the electric ray genus Diplobatis with notes on 
the interrelationships of Narcinidae (Chondrich- 
thyes, Torpediniformes).—Bulletin of the Flor- 
ida State Museum 29(5):171—209. 

Leviton, A. E., R. H. Gibbs, Jr, E. Heal, & C. E. 
Dawson. 1985. Standards in herpetology and 
ichthyology: Part I. Standard symbolic codes 


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for institutional resource collections in herpe- 
tology and ichthyology.—Copeia 1985(3):803— 
832. 

McEachran, J. D. 1995. Narcinidae. Pp. 746-792 in 
W. Fischer, EF Krupp, W. Schneider, C. Sommer, 
K. E. Carpenter and V. H. Niem, eds., Pacifico 
Centro-Oriental. Vol. II]. Rome: Organizacion 
de las Naciones Unidas para la Agricultura y la 
Alimentacion. 

Stehmann, M. 1978. Batoid fishes. in W. Fischer, ed., 
FAO identification sheets for fisheries purposes, 
Western Central Atlantic. Rome: Food and Ag- 
riculture Organization of the United Nations. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):574-578. 2001. 


A new species of “Whale Catfish” (Siluriformes: Cetopsidae) from 


the western portions of the Amazon basin 


José Carlos de Oliveira, Richard P. Vari,* and Carl J. Ferraris, Jr. 


(JCO) Universidade Federal de Juiz de Fora, Caixa Postal 656, Juiz de Fora, 

MG 36016-330, Brazil; 
(RPV) Department of Systematic Biology—Fishes, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560-0159, U.S.A.; 


(CJF) Research Associate, Department of Systematic Biology—Fishes, National Museum of 


Natural History, Smithsonian Institution, Washington, D.C. 20560-0159, U.S.A. 


Abstract.—Cetopsis parma, a new species of the subfamily Cetopsinae of 
the catfish family Cetopsidae is described from locations in the Peruvian and 
Ecuadorian Amazon. The species differs from the other species in the subfam- 
ily by the combination of the presence of a single row of conical teeth on the 
dentary, the presence of a distinct blotch of dark pigmentation on the lateral 
surface of the body dorsal to the pectoral fin, and the possession of 44 or 45 
vertebrae, 14 or 15 ribs, and 8 or 9 gill rakers. 

Resumo.—Cetopsis parma, uma nova espécie da subfamilia Cetopsinae da 
familia de bagres Cetopsidae, é descrita com base em coletas realizadas na 
Amazonia Peruana e Equatoriana. Esta espécie difere das outras espécies na 
subfamilia pela combina¢cao da presen¢a de uma Unica série de dentes cdénicos 
no dentario, da presenga de uma distinta mancha de pigmenta¢ao na superficie 
lateral do corpo dorsalmente a nadadeira peitoral, e da presenga de 44 ou 45 


vértebras, 14 ou 15 costelas, e 8 ou 9 rastros branquiais. 


The members of the Neotropical siluri- 
form subfamily Cetopsinae are commonly 
called ““Whale Catfishes” in English given 
the perceived similarity in overall form of 
cetaceans and some of the first described 
species of the subfamily. The reduced, or in 
one species absent, eyes typical of cetopsins 
are, in turn, the basis for their common 
name of ““Ciego”’ (=Blind) or ““Bagre Cie- 
go” (=Blind Catfish) in various portions of 
their range. Cetopsins have long been a 
puzzle within catfish systematics. Recently 
de Pinna & Vari (1995), however, docu- 
mented a number of unusual derived mod- 
ifications which demonstrated that the Ce- 
topsinae (the Cetopsidae of earlier authors) 
was monophyletic. The evidence indicated 
furthermore that the Cetopsinae was the sis- 


* Correspondant: vari.richard@nmnh.-si.edu. 


ter group of what had previously been rec- 
ognized as a separate family, the Helogen- 
idae. de Pinna & Vari consequently united 
the Cetopsidae and Helogenidae of previous 
classifications in an expanded Cetopsidae. 
In commenting on this broader Cetopsidae, 
de Pinna (1998:292) subsequently noted 
that “‘there is some evidence that they oc- 
cupy a markedly basal position within the 
siluriform cladogram,”’ thus making an un- 
derstanding of the species diversity and in- 
trarelationships within the family of partic- 
ular import. 

The recognized species diversity in the 
Cetopsinae has steadily increased within the 
last decade, with the 12 species of cetopsins 
considered valid by Burgess (1989) supple- 
mented by four additional species subse- 
quently described by Ferraris & Brown 
(1991), Lundberg & Rapp  Py-Daniel 


VOLUME 114, NUMBER 3 


(1994), and Ferraris (1996). Ongoing stud- 
ies indicate that these 16 species are a sub- 
stantial underestimate of the actual diversity 
in the subfamily. The species described in 
this paper is based on two specimens, one 
discovered by the first author during his ex- 
amination of the Cetopsinae (Oliveira, 
1988) and the second found by the second 
and third authors in the course of their re- 
visionary study of that subfamily. The new 
species is described herein to make the 
name available for an ongoing phylogenetic 
analysis of the Cetopsidae and a revisionary 
study of the subfamily Cetopsinae. 


Materials and Methods 


The concepts of the Cetopsidae and Ce- 
topsinae used in this paper are those pro- 
posed by de Pinna & Vari (1995). Standard 
length (SL) was measured with dial calipers 
to 1.0 mm. All measurements were taken as 
straight line distances between points. Head 
length (HL) was measured from the snout 
tip to the end of the fleshy gill cover. In- 
terorbital width was taken as the shortest 
distance between the orbits, but is difficult 
to measure unambiguously. Vertebrae and 
unpaired fin rays were counted from radio- 
graphs. Vertebral counts included the four 
elements of the Weberian complex and one 
element for the ural complex and were sep- 
arated into preanal, precaudal, and caudal 
elements. Total vertebrae is the sum of the 
precaudal and caudal vertebrae. In fin-ray 
counts, unbranched rays are indicated by 
lower case roman numerals and branched 
rays by Arabic numbers. The range of val- 
ues for meristic and morphometric features 
in the species is presented first, followed by 
the values for the holotype in brackets. In- 
stitutional abbreviations are: Museu de His- 
toria Natural de la Universidad Nacional 
Mayor de San Marcos, Lima, Peru 
(MUSM) and Museo, Escuela Politécnica 
Nacional, Quito, Ecuador (MEPN). 


Cetopsis parma, new species 
Fig. 1 
Holotype.—MUSM 2266, 73 mm SL. 
Peru. Departamento de Ucayali, Provincia 


JTS) 


Coronel Portillo, Rio Tambo, Rio Ucayali 
basin, Pucallpa, Atalaya (8°23’S, 74°32'W), 
collected by Hernan Ortega, 15 May 1986. 
Paratype.—MEPN 1034, 170 mm SL. 
Ecuador. Provincia de Pastaza, Rio Mara- 
fon basin, Rio Pastaza system, near Rio 
Chicherota, in vicinity of Montalvo (2°04'S, 
76°58'W), collected by Roman Olalla and 
Gonzalo Herrera, February 1958. 
Diagnosis.—The presence of a single 
row of teeth on the dentary in Cetopsis par- 
ma differentiates this species from all other 
cetopsins other than for Cetopsis coecutiens 
and Hemicetopsis candiru. The possession 
of conical rather than incisiform teeth dis- 
tinguishes Cetopsis parma from Hemice- 
topsis candiru and the two species also dif- 
fer in the overall form of the head and body. 
Cetopsis parma has a relatively stout body 
with the depth at the dorsal-fin origin ap- 
proximately 3.7 times in SL and the pelvic- 
fin insertion at, or slightly posterior of, the 
vertical through the posterior of the dorsal- 
fin base whereas Hemicetopsis candiru has 
an elongate body with the body depth at the 
dorsal-fin origin approximately 5 to 5.5 
times in SL and the pelvic-fin insertion dis- 
tinctly posterior of the vertical through the 
posterior of the dorsal-fin base. Cetopsis 
parma differs from C. coecutiens in its pos- 
session of the diffuse dark patch on the lat- 
eral surface of the body dorsal to the pec- 
toral fin (Fig. 1) which is lacking in the 
latter species. Cetopsis parma can be fur- 
ther differentiated from C. coecutiens in the 
total number of vertebrae (44 or 45 versus 
47 to 50, respectively), number of gill rak- 
ers (8 or 9 versus. 38 to 52, respectively), 
and number of ribs (14 or 15 versus 15 to 
18, typically 16 or 17, respectively). 
Description.—Body stout, slightly later- 
ally compressed anteriorly, increasingly 
more so posteriorly. Body depth at dorsal- 
fin origin approximately 3.7—4.2 [3.7] times 
in SL, and slightly less than HL. Lateral 
line on body complete, unbranched, mid- 
lateral, and extending from vertical through 
pectoral-fin base to hypural plate. Dorsal 
profile of body straight and obliquely slant- 


576 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 
Rio Tambo, Pucallpa, Atalaya (8°23’S, 74°32'W). 


ed from nape to dorsal-fin origin, straight 
from dorsal-fin origin to caudal-fin base. 
Ventral profile of body convex along ab- 
domen, approximately straight, but poster- 
odorsally slanted, along anal-fin base. Cau- 
dal-peduncle depth slightly greater than 
caudal-peduncle length in holotype, slightly 
greater than caudal-peduncle length in 
much larger paratype. Caudal peduncle dis- 
tinctly compressed transversely. 

Head in lateral view triangular with 
bluntly rounded snout. Dorsal profile of 
head gently convex from tip of snout to ver- 
tical through anterior margin of eye, more 
rounded from that line to nape. Ventral pro- 
file of head convex. Profile of snout in dor- 
sal view broadly rounded. Profiles of post- 
orbital portion of each side of head running 
in parallel. Dorsal surface of postorbital 
part of head with enlarged jaw musculature 
obvious. Laterosensory canals and pores on 
head not obvious. 

Branchial membranes attached to isth- 
mus posteriorly as far as vertical through 
pectoral-fin origin. Opercular opening mod- 
erate, extending ventral of horizontal 
through pectoral-fin origin for distance 
equal to snout length and dorsal of pectoral- 
fin origin for distance slightly less than 
snout length. 

Eye situated on lateral surface of head; 
located one orbital diameter dorsal of hor- 
izontal through pectoral-fin origin; eye vis- 
ible in lateral and dorsal views. Middle of 
orbit located slightly anterior to anterior 
one-quarter of HL. Eye diameter approxi- 
mately one-third length of snout in holo- 


Holotype of Cetopsis parma, new species; MUSM 2266, 73 mm SL; Peru, Departamento de Ucayali, 


type, apparently proportionally smaller but 
impossible to measure accurately in much 
larger paratype as consequence of thicker 
skin overlying eye. Interorbital width dis- 
tinctly larger than distance from tip of snout 
to rear of orbit and approximately 2.0—2.5 
[2.5] in HL. Anterior narial opening circu- 
lar, surrounded by short, anteriorly-directed, 
tubular rim of skin and located along hori- 
zontal through both tip of snout and max- 
illary-barbel origin. Distance between an- 
terior nares approximately equal to length 
of snout plus orbit. Posterior narial opening 
nearly round and without obvious long axis; 
located on dorsal surface of snout at vertical 
through anterior margin of orbit. Anterior 
two-thirds of narial opening bordered by 
flap of skin only slightly higher anteriorly. 
Distance between posterior nares slightly 
less than distance between anterior nares. 
Mouth inferior, wide, its width approxi- 
mately one-half of HL. Margin of lower 
jaw nearly transverse, its posterior limit 
reaching vertical through posterior margin 
of orbit. Premaxillary tooth patch elongate 
and crescentic, continuous across midline; 
anterior margin convex, posterior margin 
transversely aligned and nearly straight. 
Premaxillary teeth relatively small, conical, 
and sharply pointed, with teeth arranged in 
four or five irregular rows (five rows in 
larger paratype). Palatal teeth arranged in 
one gently curved row continuous across 
midline. Palatal teeth large and bluntly con- 
ical. Dentary with one row of teeth similar 
in size and shape to those on palate. 
Maxillary barbel slender, its length ap- 


VOLUME 114, NUMBER 3 


proximately equal to length of snout; barbel 
origin located along vertical through ante- 
rior margin of orbit. Mental barbels about 
equal in size and length to maxillary barbel 
and to each other. Origin of medial-mental 
barbel located at vertical through posterior 
margin of orbit. Origin of lateral-mental 
barbel located slightly posterior to vertical 
through posterior margin of orbit. Tip of 
adpressed mental barbels not reaching mar- 
gin of branchial membranes. 

Dorsal-fin rays 1,6 [1,6]. Dorsal fin rela- 
tively small, with length of base approxi- 
mately one-third of HL. Distal margin of 
dorsal fin straight, with first ray longest and 
equal in length to one-half of HL. Dorsal- 
fin spinelet absent, first dorsal-fin ray not 
spinous and with short filamentous exten- 
sion. Dorsal-fin origin located slightly pos- 
terior of one-third of SL and at vertical ex- 
tending through distal one-quarter of ad- 
pressed pectoral fin. Tip of adpressed dorsal 
fin reaching to vertical through pelvic-fin 
base. Last dorsal-fin ray without posterior 
membranous attachment to body. 

Principal caudal-fin rays 8+9 [8+9]. 
Caudal fin moderately forked and symmet- 
rical; tips of lobes bluntly pointed. Length 
of longest caudal-fin rays about one and 
one-half times length of middle rays. 

Anal-fin rays v—vi,18—25 [v,25]. Anal-fin 
base relatively short, approximately 3.7—4.2 
[3.7] times in SL. Anal-fin origin located 
well posterior to middle of SL and slightly 
posterior to middle of TL. Anal-fin margin 
straight. First branched anal-fin ray longest, 
with subsequent rays becoming gradually 
shorter. Last anal-fin ray without posterior 
membranous attachment to body. 

Pelvic-fin rays 1,5 [i,5]. Pelvic fin short, 
with distal margin slightly convex and first 
branched ray longest. Pelvic-fin origin lo- 
cated anterior to middle of SL and just pos- 
terior to vertical through posterior terminus 
of dorsal-fin base. Tip of adpressed pelvic 
fin extending beyond middle of SL, but not 
reaching vent. Last pelvic-fin ray with 
membranous attachment to body along its 
basal one-half. 


DIT 


Pectoral-fin rays 1,8—9 [1,8]. Pectoral fin 
very short, its length about one-half of HL. 
Pectoral-fin margin slightly sigmoid with 
first and middle rays longest. First pectoral- 
fin ray not spinous and with very short fil- 
amentous extension in both specimens. 

Preanal vertebrae 19—20 [19]. Precaudal 
vertebrae 15—16 [15]. Caudal vertebrae 29 
[29]. Total vertebrae 44—45 [44]. Ribs 14— 
15 [14]. Gill rakers 8—9 [9]. 

Coloration.—Head and body slightly 
darker dorsally. Sides of body dark as far 
ventrally as level of horizontal through pec- 
toral-fin base. Irregular, vertically-elongate 
dark blotch on lateral surface of body dorsal 
to pectoral fin. Height of blotch approxi- 
mately equal to length of pectoral fin, pro- 
portionally slightly higher in paratype. Ven- 
tral surface of head and abdomen pale. 
Sides of head dark as far ventrally as hor- 
izontal through base of maxillary barbel. 
Snout margin dark. Dorsal fin pale in ho- 
lotype, irregularly covered with dark pig- 
mentation in paratype. Caudal fin pale in 
holotype, covered with scattered eye-sized 
dark spots in much larger paratype. Anal fin 
dusky basally and pale distally in both spec- 
imens. Pectoral fin with interradial mem- 
branes darkly pigmented on dorsal surface 
except along fin margin. Pelvic fin with 
scattered dark pigmentation on dorsal sur- 
face of interradial membranes, except along 
fin margin. 

Sexual dimorphism.—Both the holotype 
and paratype are presumed to be females, 
given their possession of the straight anal- 
fin margin which is typical of females in 
the sexually dimorphic species in the Ce- 
topsinae (RPV & CJF pers. obs). 

Distribution.—Cetopsis parma is only 
known from two localities in the western 
portions of the Amazon basin in the Rio 
Pastaza of the Rio Marafion basin in Ec- 
uador and the Rio Tambo in the Rio Ucayali 
basin of Peru. 

Etymology.—The species name, parma, 
is from the Latin word, parma, a type of 
small shield, is in reference to the dark 
mark on the lateral surface of the body im- 


578 


mediately dorsal to the pectoral fin. It is 
used as a noun in apposition. 

Remarks.—Various authors (Ferraris & 
Brown 1991, Lundberg & Rapp Py-Daniel 
1994, and Ferraris 1996) have grappled 
with the problem of generic definitions 
within the subfamily Cetopsinae (the family 
Cetopsidae of authors prior to de Pinna & 
Vari 1995). Lundberg & Rapp Py-Daniel 
(1994:381) well summarized the situation 
with their comment that the ““systematic un- 
derstanding of the South American Cetop- 
sidae is poorly developed’’. The resolution 
of these problems lies far beyond the limits 
of this study and will be addressed in a fu- 
ture publication dealing with the phyloge- 
netic relationships among cetopsins. For the 
purposes of this paper, we consequently 
place the new species into Cetopsis using 
the concept of that genus proposed by 
Schultz (1944) which is based on the com- 
bination of the possession of a restricted gill 
opening and conical teeth arranged in mul- 
tiple rows on the premaxillae. We recog- 
nize, however, that the limits of the genus 
may be modified by a more rigorous phy- 
logenetic analysis. 

The holotype and paratype of Cetopsis 
parma demonstrate various morphometric 
differences which likely reflect the pro- 
nounced differences in the size of the spec- 
imens (73 versus 170 mm SL, respectively). 
The holotype and paratype show notewor- 
thy variation in only one meristic feature, 
the number of branched anal-fin rays (18 
versus 25, respectively) and the associated 
morphometric value, the proportional 
length of the base of the anal fin. Until such 
time as additional material which shares the 
distinctive pigmentation and dentition char- 
acters present in these specimens becomes 
available, we conservatively consider this 
difference to represent intraspecific varia- 
tion, perhaps reflective of populational dif- 
ferences associated with the river distances 
separating the two localities. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Acknowledgments 


Research and museum visits associated 
with this study were made possible by fund- 
ing through the Neotropical Lowlands Re- 
search Program of the Smithsonian Institu- 
tion. We thank Hernan Ortega (MUSM) and 
Ramiro Barriga (MEPN) for the loan of 
specimens which made this paper possible. 
Sandra J. Raredon provided technical sup- 
port for the project. Fig. 1 was prepared by 
T. Britt Griswold. This paper benefitted 
from comments and suggestions from Mar- 
io C.C. de Pinna, Scott A. Schaefer, and an 
anonymous reviewer. 


Literature Cited 


Burgess, W. E. 1989. An atlas of freshwater and ma- 
rine catfishes. A preliminary survey of the Sil- 
uriformes. T.-FH. Publications, Neptune City, 
New Jersey, U.S.A. 784 pp. 

Ferraris, C. J., Jr. 1996. Denticetopsis, a new genus of 
South American whale catfish (Siluriformes: 
Cetopsidae, Cetopsinae), with two new spe- 
cies.—Proceedings of the California Academy 
of Sciences 49(6):161—170. 

, & B. A. Brown. 1991. A new species of Pseu- 
docetopsis from the Rio Negro drainage of Ven- 
ezuela (Siluriformes: Cetopsidae).—Copeia 
1991(1):161—165. 

Lundberg, J. G., & L. Rapp Py-Daniel. 1994. Bathy- 
cetopsis oliveirai, gen. et sp. nov., a blind and 
depigmented catfish (Siluriformes: Cetopsidae) 
from the Brazilian Amazon.—Copeia 1994(2): 
381-390. 

Oliveira, J. C. de 1988. Osteologia e revisao sistema- 
tica de Cetopsidae (Teleostei, Siluriformes) Un- 
published Ph.D. dissertation, Universidade de 
Sao Paulo. 

de Pinna, M. C. C. 1998. Phylogenetic relationships of 
Neotropical Siluriformes (Teleostei: Ostario- 
physi): Historical overview and synthesis of hy- 
potheses. Pp. 279-330 in L. R. Malabarba, R. 
E. Reis, R. P Vari, Z. M. S. Lucena, and C. A. 
S. Lucena, eds., Phylogeny and Classification of 
Neotropical Fishes. Porto Alegre, Brazil, Edi- 
pucrs, 603 pp. 

, & R. P Vari. 1995. Monophyly and phylo- 
genetic diagnosis of the family Cetopsidae, with 
synonymization of the Helogenidae (Teleostei: 
Siluriformes).—Smithsonian Contributions to 
Zoology 51:1—26. 

Schultz, L. PB 1944. The catfishes of Venezuela, with 
descriptions of thirty-eight new forms.—Pro- 
ceedings of the United States National Museum 
94 (3172):173-338, pls. 1-14. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):579—588. 2001. 


A new species of the luvarid fish genus {Avitoluvarus (Acanthuroidei: 
Perciformes) from the Eocene of the Caucasus in southwest Russia 


Alexandre EK Bannikov and James C. Tyler 


(AFB) Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, 
Moscow 117647, Russia; 
(JCT) National Museum of Natural History, Smithsonian Institution, 
Washington, D.C. 20560-0106, U.S.A. 


Abstract.—A new species of the luvarid fish genus fAvitoluvarus, A. eocaen- 
icus, 1S described from the middle Eocene of Russia (Kuma Horizon, North 
Caucasus) based on a single imprint of its skeleton (incomplete posteriorly). 
Avitoluvarus previously has been known only from the terminal (uppermost) 
Paleocene of Turkmenistan, where it is represented by two species, A. dianae 


and A. mariannae. 


The fossil luvarid genus Avitoluvarus 
was described recently (Bannikov & Tyler 
1995) from the Danata Formation in Turk- 
menistan, where it is represented by two 
species, A. dianae Bannikov & Tyler, 1995, 
and A. mariannae Bannikov & Tyler, 1995. 
The exact age of the fish-bearing layer of 
the Danata Formation in Turkmenistan has 
been questioned, with the fishes from this 
layer dated as both Late Paleocene (Danil- 
chenko 1968, Bannikov 1985) and Early 
Eocene (Tyler & Bannikov 1992b, Patter- 
son 1993, Bannikov & Tyler 1995). We ac- 
cept here the analysis of Muzylev (1994) 
indicating that the fish-bearing layer of the 
Danata Formation is synchronous with the 
Upper Thanetian sapropel of more western 
regions, corresponding to global Late Pa- 
leocene anoxic events. 

In 1999, excavations in the North Cau- 
casus by the first listed author yielded the 
imprint of a skeleton (incomplete posteri- 
orly) that represents a new species of Avi- 
toluvarus. This specimen was found in the 
bituminous marls of the Kuma Horizon in 
the Gorny Luch locality (Pshekha River, 
Apsheronsk District, about 0.5 km from the 
Gorny Luch farmstead). The Kuma Horizon 
correlates with the Bartonian stage. The 
Kuma Horizon previously has been as- 


signed to the Upper Eocene (Tyler & Ban- 
nikov 1992a, Bannikov 1993) but according 
to Cavelier & Pomerol (1986) only the Pria- 
bonian (but not the Bartonian) should be 
included in the Upper Eocene, and thus the 
Kuma Horizon fishes are of late Middle Eo- 
cene age. 

A preliminary, and far from complete, 
list of Kuma fishes (Bannikov & Parin 
1997) includes at least 27 species repre- 
senting 25 families in 10 orders. Based on 
the recent excavations in 1999, this list can 
be extended by a number of taxa repre- 
senting their first discoveries at the generic 
and familial levels at the Gorny Luch lo- 
cality; e.g., a champsodontid, an Antigonia- 
like caproid, a Seriola-like carangid, a Sar- 
da-like scombrid, a percoid fish probably 
belonging to a new genus of uncertain fam- 
ily, and the new species of luvarid de- 
scribed below. The great majority of the tel- 
eost species from the Gorny Luch assem- 
blage are oceanic pelagic (epi- and meso-) 
and the new species of Avitoluvarus de- 
scribed below is presumably epipelagic like 
the only Recent representative of the fam- 
ily, Luvarus imperialis Rafinesque, 1810. 

The new species represents the first re- 
cord of the genus Avitoluvarus outside of 
Turkmenistan and extends the stratigraphic 


580 


distribution of this genus from the Late Pa- 
leocene to the end of the Middle Eocene. 


Family Luvaridae Gill, 1885 
Genus Avitoluvarus Bannikov & Tyler, 
1995 
Avitoluvarus eocaenicus, new species 
Figs. 1-3 


Material.—Holotype, Paleontological In- 
stitute (PIN), Moscow, no. 4425/32, esti- 
mated 38 mm SL, imprint of poorly pre- 
served skeleton, incomplete posteriorly, 
single plate, and counterpart of the head 
and cleithrum. 

Type locality and horizon.—Left bank of 
Pshekha River, about 0.5 km from the Gor- 
ny Luch farmstead, Apsheronsk District, 
Krasnodar Region; Kuma Horizon, upper 
part of the Middle Eocene. 

Etymology.—For the Eocene age of the 
new species. 

Diagnosis.—Avitoluvarus eocaenicus 
has all the available characters (those of the 
caudal peduncle are unknown because of 
the incompleteness posteriorly of the holo- 
type) diagnostic of the genus Avitoluvarus 
(see diagnosis in Bannikov & Tyler 1995: 
6). Avitoluvarus eocaenicus differs from 
both of the other two species of the genus, 
A. dianae and A. mariannae, by the anter- 
oventral inclination of the third to fifth hae- 
mal spines, versus these spines being ori- 
ented relatively vertically in A. dianae and 
inclined posteroventrally in A. mariannae 
(for comparative data on the two previously 
described species, here and following, see 
Bannikov & Tyler 1995). Additionally, A. 
eocaenicus differs from the other two spe- 
cies by a combination character: the haemal 
spines of the first two caudal vertebrae are 
slender and make contact with one another 
in their middle regions, versus these haemal 
spines being slender but not contacting one 
another in A. mariannae, or relatively stout 
and contacting one another in A. dianae. 
These diagnostic haemal spine characters 
do not change appreciably ontogenetically 
within the size ranges known for the other 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


two species of Avitoluvarus, and the single 
specimen of A. eocaenicus (38 mm SL) is 
within the size range of the specimens of 
A. mariannae (34-190 mm SL) and A. 
eocaenicus (38-120 mm SL). 

Description.—The holotype of Avitolu- 
varus eocaenicus is probably a juvenile 
based on the presence of ridges on the skull 
bones and serrations on the dorsal- and pel- 
vic-fin spines. Such ridges and serrations 
are present in other small juvenile (less than 
about 50 mm SL) luvarids (e.g., as in A. 
mariannae and Luvarus imperialis) but 
these are lost at larger sizes. Also, the skel- 
eton of A. eocaenicus seems to be about as 
weakly ossified as in A. mariannae and 
somewhat less well ossified than in A. di- 
andae. 

The body is fusiform, and its greatest 
depth is about 41% SL. The head is rela- 
tively long, about 35% SL. The upper head 
profile is gently curved and the mouth is 
small (gap equal to about orbit diameter or 
less). The round orbit is situated in the low- 
er part of the upper half of the head and has 
a horizontal diameter of about 23% of the 
head length. 

Skull: The limits of the individual bones 
of the weakly ossified occipital and otic re- 
gions are not clear, but the frontals seem to 
be relatively wide and the supraoccipital 
lacks a prominent crest. The bones of the 
cranial roof (probably of the frontals) have 
remnants of larval ridges, and we presume 
that these ridges were serrate as in other 
acanthuroids even though we cannot be ab- 
solutely sure of this because of poor pres- 
ervation of the single specimen; one ridge 
is along the dorsal midline and another, 
shorter one, is anterior to the orbit. 

The ethmoid region is exceptionally 
weakly ossified. The parasphenoid is slen- 
der and slightly convex where it is exposed 
at the lower edge of the orbit. The shaft of 
the hyomandibular is oriented anteroven- 
trally. The pterygoid bones and the palatine 
are unclear. The quadrate is broad, trian- 
gular, and has a small articular facet for the 
lower jaw. The symplectic is an elongate 


VOLUME 114, NUMBER 3 


581 


Fig. 1. 
SL Gncomplete posteriorly), wet with alcohol to improve contrast, Middle Eocene of North Caucasus (Kuma 
Horizon), Russia. Scale bar is 10 mm. 


rod, somewhat stouter posteriorly than an- 
teriorly. There is a faint remnant of a struc- 
ture in the ethmoid region above the ante- 
rior part of the parasphenoid that can be 
interpreted as a long slender process or 
prong of the lachrymal similar to that of 
juvenile specimens of Luvarus imperialis 
(see Tyler et al. 1989). 

The lower jaw articulation is situated 
well in front of the level of the anterior 
edge of the orbit. The lower jaw is rather 
short but deep, with the dentary and artic- 
ular of about equal size. The dentary bears 
a single row of small conical teeth. The al- 
veolar process of the premaxilla is slender, 
elongate, and bears a single row of small 
conical teeth, whereas the ascending pro- 
cess is short but prominent. The maxilla 
seems to be relatively wide. 

The thin, flat bones of the opercular re- 
gion are poorly preserved, with the limits 
of the individual bones unclear. The opercle 
appears to be subtriangular and has several 
bony ridges radiating from the condylar re- 
gion. The preopercle is curved along its an- 
terior border, with an angle of about 100°. 


Avitoluvarus eocaenicus, new species, photograph of the holotype, PIN 4425/32, estimated 38 mm 


Several radiating ridges appear to be pre- 
sent in the upper part of the preopercle. 

The hyoid and branchial arches are not 
clear, but there are five sabre-like bran- 
chiostegal rays. 

Pectoral fin and girdle: The posttemporal 
is slender and elongate; it extends from the 
posterodorsal aspect of the head ventrally 
and slightly posteriorly. A short intercalar 
process extends anteriorly from the lower 
part of the posttemporal. The supracleith- 
rum is elongate and tapered anterodorsally; 
this bone is disarticulated in the present 
specimen. The large cleithrum has a gently 
curved c-shape, with the upper and lower 
ends inclined, respectively, slightly antero- 
dorsally and anteroventrally. The upper lim- 
its of the postcleithrum are poorly pre- 
served, but the long shaft of the bone below 
the pectoral-fin base is sturdy and reaches 
ventrally below the ventral edge of the an- 
terior extension of the first anal-fin ptery- 
giophore (this extension is probably dis- 
placed postmortem somewhat dorsally) or 
almost to the ventral margin of the body. 
There is no evidence of a division of the 


582 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Avitoluvarus eocaenicus, new species, reconstruction of the holotype (for data see legend for Fig. 
1). The unstippled parts of the skeleton posterior to the dashed fracture line through the sixth caudal vertebra 
are missing and hypothesized here as having the conditions of all other species of the family Luvaridae (e.g., a 
total of 13 caudal vertebrae, the pterygial truss extending posteriorly to the ninth caudal vertebra, four vertebrae 
in the caudal peduncle, and a forked caudal fin deeply overlapping the hypural plate). 


VOLUME 114, NUMBER 3 


long postcleithral shaft into two pieces, and 
we presume that the postcleithrum is a sin- 
gle bone (the right and left postcleithra are 
preserved close alongside one another, ex- 
cept where they overlap ventrally, and are 
so shown in the reconstruction). The cora- 
coid is elongate, of decreasing width ante- 
roventrally. 

The base of the pectoral fin is situated in 
the middle of the body, slightly above the 
middle of the distance between the verte- 
bral column and the ventral profile. The 
pectoral fin is poorly preserved and its 
length and number of rays cannot be deter- 
mined. 

Pelvic fin and girdle: The pelvis is L- 
shaped. The long ascending (pubic) process 
is oriented slightly posterodorsally toward 
the cleithrum. The posterior (ishial) process 
is well developed, reaching almost to the 
level of the anterior end of the anterior ex- 
tension of the first anal-fin pterygiophore. 
The posterior process is tapered and shorter 
than the ascending process. There is essen- 
tially no anterior (iliac) process. 

The pelvic spine is relatively long (lon- 
ger than the posterior process of the pelvis), 
about 8% SL, robust, and bears serrations 
along its anterior edge, being similar to the 
first dorsal-fin spine except somewhat 
shorter. No soft rays are evident in the pel- 
vic fin. The pelvic fin of luvaroids becomes 
rudimentary with increasing specimen size, 
or it is absent (Bannikov & Tyler 1995:33), 
so it can be anticipated that when larger 
specimens of A. eocaenicus become avail- 
able the pelvic-fin spines will be much 
shorter (or absent) than in the present spec- 
imen of ca. 38 mm SL. 

Vertebral column: There are probably 9 
+ 13 = 22 vertebrae, as in all other species 
of both fossil and Recent luvarids; if so, the 
posterior seven vertebrae are missing in the 
only specimen of A. eocaenicus (the miss- 
ing vertebrae are indicated hypothetically 
by dashed lines in the reconstruction, Fig. 
2). The first two abdominal vertebrae are 
obscured, but can be reasonably estimated 
on the basis of the space available for them 


583 


between the rear of the skull and the front 
of the centrum that is the first to be clearly 
exposed (presumably the third abdominal 
vertebra). The vertebral column is elevated 
anteriorly and articulates relatively high on 
the rear of the cranium. All of the preserved 
centra are somewhat elongate anteroposte- 
riorly. All of the preserved neural spines are 
short, slender, and only slightly curved. 
Most of the neural spines are inclined pos- 
teriorly, but the orientation of those of the 
last abdominal and first two caudal verte- 
brae is close to vertical. The neural spines 
of the first five caudal vertebrae are shorter 
than the corresponding haemal spines. The 
abdominal vertebrae lack parapophyses. 
Short and slender ribs (pleurals) are present 
on the third to ninth abdominal vertebrae, 
becoming slightly shorter posteriorly. The 
ribs are inclined posteroventrally and reach 
to a level less than one-half the distance 
between the vertebral column and the ven- 
tral profile of the body. Epineurals are not 
evident. 

The haemal spine of the first caudal ver- 
tebra is only slightly thicker than that of the 
second caudal vertebra; it is only moderate- 
ly curved anteroventrally in its lower re- 
gion. The first haemal spine is so closely 
articulated along its anterior edge to the first 
anal-fin pterygiophore that its full length 
cannot be determined. The haemal spine of 
the second caudal vertebra is very slender 
and almost straight. The second haemal 
spine is angled anteriorly from its base at 
the centrum and closely approaches or con- 
tacts the posterior edge of the middle of the 
first haemal spine, distal to which point of 
contact these two haemal spines diverge. 
The haemal spines of the third to sixth cau- 
dal vertebrae are exceptionally slender, 
straight or only slightly curved, and of de- 
creasing length posteriorly in the series. 
The third to fifth haemal spines are inclined 
anteroventrally, whereas the sixth haemal 
spine is oriented slightly posteroventrally, 
and close to vertical. 

The caudal peduncle, caudal skeleton, 
and caudal fin are missing. 


584 


Dorsal and anal fins: The posterior parts 
of these fins are missing and the total num- 
ber of dorsal- and anal-fin elements cannot 
be determined exactly. However, based on 
the assumption that the missing parts of the 
dorsal fin are similar to those of other lu- 
varids, we estimate that there are about 24 
dorsal-fin elements. The first dorsal-fin el- 
ement is definitely a spine. It is relatively 
long, about 11% SL, borne in supernumer- 
ary association on the first dorsal-fin pter- 
ygiophore, and bears serrations along its an- 
terior edge. The second and third elements 
are represented by faint remnants; these are 
evidently thicker than the subsequent ele- 
ments, and therefore are probably spines. 
The remaining elements are clearly soft 
rays that are unsegmented, unbranched, and 
bilaterally paired (as seen in some elements 
that are separated into slightly displaced left 
and right halves). The rays are much shorter 
than the first dorsal-fin spine. The dorsal- 
fin pterygiophores are very poorly pre- 
served and the majority of their ventral 
shafts are unclear. The distal expansions of 
the dorsal-fin pterygiophores apparently 
form a continuous truss, but we cannot de- 
termine its thickness. The upper region of 
the ventral shaft of the first dorsal-fin pter- 
ygiophore is expanded into lamellar plates 
both anteriorly and posteriorly. Based on its 
position, the lower region of the ventral 
shaft of the first dorsal-fin pterygiophore is 
situated in what we estimate to be the pre- 
neural space in front of the distal end of the 
neural spine of the first vertebra. 

Only a few of the anal-fin rays are pre- 
served, in about the middle of the fin, but 
based on the number of pterygiophores pre- 
served and of missing parts being similar to 
those of other luvarids, we estimate that 
there are about 22 anal-fin rays. The anal- 
fin rays are unsegmented, unbranched, and 
bilaterally paired, similar to those of the 
dorsal fin and of about the same length. The 
anal-fin pterygiophores are mostly T-shaped 
in lateral view, with a dorsally oriented 
shaft and an anteroposteriorly expanded 
distal end. The first anal-fin pterygiophore 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


has a long, thick anterior extension beneath 
most of the abdominal cavity, and a long, 
equally stout posterodorsal process whose 
upper end firmly articulates along the an- 
terior edge of the lower end of the haemal 
spine of the first caudal vertebra. The sec- 
ond and more posterior anal-fin pterygio- 
phores have the distal ends expanded into 
shallow anterior and posterior processes 
that form a continuous truss that we pre- 
sume must have been extensively interdig- 
itated even though we cannot determine the 
details of this because of poor preservation. 

As preserved, the ventral edge of the 
truss in the abdominal region just behind 
the anterior extension of the first anal-fin 
pterygiophore is strongly convex (Figs. 1, 
2), uniquely so among luvarids. However, 
we do not believe that this is an autapo- 
morphy of the new species, but, rather, a 
postmortem artifact associated with the up- 
ward displacement of the anterior extension 
of the first anal-fin pterygiophore above the 
level of the lower end of the postcleithrum; 
the weak ossification in this juvenile spec- 
imen would lend itself to this distortion. In 
Figure 3 we show this region of the truss 
as preserved (A) and as we believe it would 
appear if undistorted (B). 

With the exception of the first anal-fin 
pterygiophore, the other pterygiophores are 
very slender. The shafts of the second to 
sixth pterygiophores are situated between 
the haemal spines of the first and second 
caudal vertebrae (first interhaemal space), 
and those of the seventh to ninth pterygiop- 
hores are in the second interhaemal space. 
The succeeding three interhaemal spaces 
accommodate two or three pterygiophore 
shafts each. The shafts of the anal-fin pter- 
ygiophores are not especially convergent 
toward the haemal spines. 

Scales: Tiny scales cover the entire body, 
but poor preservation precludes detailed de- 
scription. 


Discussion 


Bannikov & Tyler (1995) proposed nine 
unequivocal synapomorphies for the super- 


VOLUME 114, NUMBER 3 


585 


Fig. 3. 


Avitoluvarus eocaenicus, new species: A, structure of the anterior pterygiophores of the anal fin as 


preserved with postulated convex distortion of the lower edge of the first pterygiophore in the region behind 
the anterior process, and with upward displacement of the anterior process above the level of the ventral end of 
the postcleithrum; B, our interpretation of how the anterior pterygiophores of the anal-fin would appear if 


undistorted. 


family Luvaroidea (Luvaridae + +Kushlu- 
kiidae) and six unequivocal synapomor- 
phies for the family Luvaridae (with two 
species of Paleocene Avitoluvarus, and two 
species of Luvarus, one Paleocene and one 
Recent; Tyler et al. 1989, previously doc- 
umented a far larger number of synapo- 
morphies for the Luvaridae based only on 
the single Recent species, but many of these 
features cannot be determined in fossils or 
are unique to the Recent species). Within 
the Luvaridae, the preponderance of de- 
rived features were found in Luvarus, 
whereas Avitoluvarus was shown to possess 
only two unequivocal synapomorphies: (1) 
the truss formed by the interdigitation of the 
distal regions of the dorsal- and anal-fin 
pterygiophores is relatively shallow and not 
extensively interdigitated; and (2) the prox- 


imal shafts of a total cf seven to 11 (rarely 
six) anal-fin pterygiophores are situated in 
the first two interhaemal spaces (here and 
following, character numbers are in paren- 
theses and correspond to those in the clad- 
ogram, Fig. 4). 

Within Avitoluvarus, Bannikov & Tyler 
(1995) identified one autapomorphy of A. 
dianae as (3) the posterodorsally oriented 
proximal shaft of the first anal-fin ptery- 
giophore is especially thick and stout, and 
three autapomorphies of a. mariannae as: 
(4) the proximal shafts of a total of 10 to 
11 anal-fin pterygiophores are situated in 
the first two interhaemal spaces (the greater 
increase in number of pterygiophores in 
these two spaces in A. mariannae being a 
more derived condition than the lesser in- 
crease in A. dianae); (5) the ribs are rela- 


586 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


KUSHLUKIA 
A. EOCAENICUS 


A. DIANAE 


A. MARIANNAE 
LUVARUS 


Fig. 4. Cladogram of the proposed relationships within Avitoluvarus. Derived character states, discussed in 
the text, are: (1) pterygial truss shallow and not extensively interdigitated; (2) proximal shafts of a total of 7— 
11 (arely 6) anal-fin pterygiophores situated in the first two interhaemal spaces; (3) posterodorsally oriented 
proximal shaft of the first anal-fin pterygiophore especially thick and stout; (4) proximal shafts of a total of 10— 
11 anal-fin pterygiophores situated in the first two interhaemal spaces (5) ribs relatively short and thin, ending 
at the level of the upper third of the abdominal cavity; (6) ventral half of pterygial truss especially shallow; (7) 
first two haemal spines only slightly thickened; (8) proximal shafts of the second and many of the more posterior 
anal-fin pterygiophores exceptionally slender; (9) third to fifth haemal spines inclined anteroventrally. Character 
states 1-6 and 9 are unequivocal apomorphies, but states 7—8 are equivocal and apomorphic only under the 
assumption of delayed transformation, favoring independent acquisition over reversal. For data supporting this 
and the sister group relationships of Kushlukia with Avitoluvarus+ Luvarus, see Bannikoy & Tyler (1995). 


tively short and thin, ending at the level of 
about the upper third of the abdominal cav- 
ity; and (6) the ventral half of the pterygial 
truss formed from the distal ends of the 
anal-fin pterygiophores is very shallow. 


Moreover, A. mariannae was shown to pos- 
sess two equivocal autapomorphies (which 
assume delayed transformation, favoring in- 
dependent acquisition over reversal, with 
the ancestor possessing the plesiomorphic 


VOLUME 114, NUMBER 3 


condition of the outgroups): (7) the first two 
haemal spines are only slightly thickened; 
and (8) the proximal shafts of the second 
and more posterior anal-fin pterygiophores, 
except for the last few, are exceptionally 
slender. 

The new species, A. eocaenicus, shares 
four derived character states (S—6 unequiv- 
ocal, 7-8 equivocal) with A. mariannae and 
lacks the single autapomorphy (3) of A. di- 
anae, all of which indicates a sister group 
relationship between A. eocaenicus and A. 
mariannae. 

For one character (4), A. eocaenicus is 
intermediate between A. dianae and A. mar- 
iannae. Two to five anal-fin pterygiophore 
shafts in the first two interhaemal spaces is 
ancestral for acanthuroids (Bannikov & Ty- 
ler 1995), and the increase in number of 
such pterygiophores in Avitoluvarus can be 
considered an ordered transformation series 
of increasing specialization from the six to 
seven in A. dianae to the eight in A. eocaen- 
icus, to the 10-11 in A. mariannae. For this 
character within Avitoluvarus, we consider 
the eight or more pterygiophores in these 
two spaces as a derived condition synapo- 
morphic for A. dianae and A. mariannae. 

We identify an autapomorphy of A. 
eocaenicus as (9) the third to fifth haemal 
spines are inclined anteroventrally. We con- 
sider posteroventral to vertical inclination 
of the third to fifth haemal spines as ances- 
tral for acanthuroids because this is the con- 
dition found in fossil and Recent scato- 
phagids (the first outgroup for acanthu- 
roids), siganids, zanclids, and most acan- 
thurids (including those considered to be 
the most basal): see illustrations in Tyler et 
al. 1989, Blot & Tyler 1991, Tyler & Ban- 
nikov 1997, Tyler & Sorbini 1999). Among 
the luvarid + +kushlukiid clade (Bannikov 
& Tyler 1995), anteroventral orientation of 
the third to fifth haemal spines is found 
only in one of the two species of Kushlukia 
(in K. permira, versus posteroventral ori- 
entation in K. sp.), in one of the two species 
of Luvarus (in L. imperialis, versus vertical 
to posteroventral in L. necopinatus), and in 


587 


one of the three species of Avitoluvarus (in 
A. eocaenicus, versus posteroventral in A. 
mariannae and vertical in A. dianae). It is 
most parsimonious to propose that the an- 
teroventral orientation has arisen indepen- 
dently in the single species of Kushlukia, 
Luvarus, and Avitoluvarus; therefore, the 
anteroventral orientation of the third to fifth 
haemal spines is an autapomorphy of A. 
eocaenicus. 


Conclusion 


The addition of another new species to 
the record of fossil luvarids reinforces the 
notion (Bannikov & Tyler 1995:40) that the 
luvaroid fishes were far more diversified in 
the early Tertiary (six species in two fami- 
lies, with four species from the uppermost 
Paleocene of the Danata Formation, Turk- 
menistan, alone) than they are today, when 
only a single species survives. 


Acknowledgments 


We thank the National Geographic So- 
ciety, Washington DC, for grant no. 6555- 
99 that allowed the first listed author to 
make excavations in the North Caucasus in 
1999, where the holotype of Avitoluvarus 
eocaenicus was obtained. The manuscript 
was improved by the suggestions of G.D. 
Johnson, Smithsonian Institution, and an 
anonymous reviewer. The photograph of 
the holotype was made by A.V. Mazin 
(PIN, Moscow). 


Literature Cited 


Bannikov, A. E 1985. Iskopayemye skumbriyevye 
SSSR [Fossil scombrids of the USSR].—Trudy 
Paleontologicheskogo Instituta Akademii Nauk 
SSSR 210:1-111. 

. 1993. The succession of the Tethys fish as- 

semblages exemplified by the Eocene localities 

of the southern part of the former USSR.—Kau- 
pia, Darmstadter Beitrage zur Naturgeschichte 

2:241—246. 

, & N.N. Parin. 1997. The list of marine fishes 

from Cenozoic (Upper Paleocene-Middle Mio- 

cene) localities in southern European Russia 
and adjacent countries.—Journal of Ichthyology 

37:133-146. 


588 


, & J.C. Tyler. 1995. Phylogenetic revision of 
the fish families Luvaridae and +Kushlukiidae 
(Acanthuroidei), with a new genus and two new 
species of Eocene luvarids.—Smithsonian Con- 
tributions to Paleobiology 81:1—45. 

Blot, J., & J. C. Tyler. 1991. New genera and species 
of fossil surgeon fishes and their relatives 
(Acanthuroidei, Teleostei) from the Eocene of 
Monte Bolca, Italy, with application of the Blot 
formula to both fossil and Recent forms.—Mis- 
cellanea Paleontologica, Studi e Ricerche sui 
Giacimenti Tertziari di Bolca, Museo Civico di 
Storia Naturale di Verona 6:13—92. 

Cavelier, C., & C. Pomerol. 1986. Stratigraphy of the 
Paleocene.—Bulletin de la Société de la Géo- 
logique de France 2(2):255—265. 

Danilchenko, P. G. 1968. Ryby verkhnego paleotsena 
Turkmenii [Fishes of the Upper Paleocene of 
Turkmenistan]. Pp. 113—156 in D. V. Obruchev, 
ed., Ocherki po filogenii i sistematike iskopay- 
emykh ryb 1 beschelyustnykh [Outlines on the 
phylogeny and systematics of fossil fishes and 
agnathans]. Nauka, Moscow, 173 pp. 

Muzylev, N. G. 1994. Anoksicheskiye sobytiya pa- 
leotsena-srednego eotsena [Anoxic events of the 
Paleocene-Middle Eocene]. Pp. 160-166 in 
A.Y. Rozanov and M.A. Semikhatov, eds., Eko- 
sistemnye perestroiki 1 evolyutsiya biosfery. 
Vypusk | [Ecosystem restructuring and the evo- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


lution of the biosphere. Issue 1] Nedra, Mos- 
cow, 367 pp. 

Patterson, C. 1993. Osteichthyes: Teleostei. Pp. 621— 
656 in M. J. Benton, ed., The fossil record 2. 
Chapman and Hall, London, 845 pp. 

Tyler, J. C., & A. E Bannikov. 1992a. New genus of 
primitive ocean sunfish with separate premax- 
illae from the Eocene of southwest Russia 
(Molidae, Tetraodontiformes).—Copeia 1992 
(4):1014—1023. 

. 1992b. A remarkable new genus of tetraodon- 

tiform fish with features of both balistids and 

ostracids from the Eocene of Turkmenistan.— 

Smithsonian Contributions to Paleobiology 72: 

1-14. 

. 1997. Relationships of the fossil and Recent 

genera of rabbitfishes (Acanthuroidei: Sigani- 

dae).—Smithsonian Contributions to Paleobi- 

ology 84:1—35. 

, & C. Sorbini. 1999. Phylogeny of the fossil 

and Recent genera of fishes of the family Sca- 

tophagidae (Squamipinnes).—Bollettino dei 

Museo Civico di Storia Naturale di Verona 23: 

353-393. 

, G. D. Johnson, I. Nakamura, & B. B. Collette. 

1989. Morphology of Luvarus imperialis (Lu- 

varidae), with a phylogenetic analysis of the 

Acanthuroidei (Pisces).—Smithsonian Contri- 

butions to Zoology 485:1—78. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):589—-598. 2001. 


Pseudothyone levini, a new species of sea cucumber (Echinodermata: 


Holothuroidea) from the northeastern Pacific 


Philip Lambert and Kathryn L. Oliver 


(PL) Natural History Section, Royal British Columbia Museum, 675 Belleville Street, 
Victoria, B.C., Canada, V8W 9W2; 
(KLO) 403-1631 McKenzie Avenue, Victoria, B.C., Canada, V8N 5M3 


Abstract.—Pseudothyone levini, new species (Sclerodactylidae), based on 12 
specimens collected between Graham Island, Queen Charlotte Islands, British 
Columbia and Tacoma, Puget Sound, Washington, U.S.A. in the northeastern 
Pacific Ocean, is described. It ranges in depth from the intertidal zone to 70 
meters on rock and in mud or gravel. Body is beige or white. Podia are scat- 
tered but more numerous in the five ambulacra. Usually ten brown dendritic 
tentacles. P. levini has mostly round or oval skin ossicles with a smooth sur- 
face, a wavy margin and small holes. Another less numerous plate is larger 
and thicker with scalloped margins and numerous large holes. Tails of the 
radials on the calcareous ring are short and curved. Tentacle ossicles range 
from large robust, curved rods perforated at the ends, to small curved, perfo- 
rated plates. Pseudothyone levini is compared to seven other species currently 


recognized in the genus. 


The family Sclerodactylidae is well rep- 
resented in the Northern hemisphere, pri- 
marily in the Atlantic, from shallow water 
to bathyal depths (Pawson 1982). Three 
species in two genera of Sclerodactylidae 
inhabit the northwestern coast of North 
America. Eupentacta pseudoquinquesemita 
Deichmann ranges from Alaska to Wash- 
ington, and EF. quinquesemita (Selenka) 
ranges from southern Alaska to southern 
California (Lambert 1997). Pachythyone 
rubra (H. L. Clark) is known from Mon- 
terey Bay to Los Angeles Harbor, Califor- 
nia (Bergen 1996). 

Since 1986, five new species of holothu- 
roids have been added to the known fauna 
of the northeastern Pacific (Lambert 1997, 
1998). Here we describe a sixth new spe- 
cies of sea cucumber based on material 
from British Columbia and Washington. 
The genus Pseudothyone is represented by 
three species in the Atlantic, one in the 
Mediterranean, one in the Persian Gulf, one 


in the western Pacific, and now this one in 
the northeastern Pacific. 


Materials and Methods 


Ossicle slides were prepared as in Lam- 
bert (1985) and measurements of ossicles 
followed the procedures described in Lam- 
bert (1998). Ossicles were sampled from 
the dorsal tentacles of all specimens, mid- 
dorsal regions of intact specimens, and 
from anterior and posterior regions of par- 
tial specimens. 


Results 


Order Dendrochirotida Grube 
Family Sclerodactylidae Pawson & Fell, 
1965 


Diagnosis.—Body lacks a test of imbri- 
cating plates; body wall soft; ossicles small 
and inconspicuous. Calcareous ring com- 
plex, with paired or unpaired processes; el- 
ements of ring not composed of a mosaic 


590 


of minute pieces. Tentacles 10—20. Body 
may be U-shaped. Calcareous ring not as 
massive as in the Phyllophoridae. Tube feet 
usually scattered in the radii and interradii, 
but tendency for them to be restricted to the 
radii (Pawson 1982). 


Subfamily Sclerodactylinae Panning, 
1949; restricted by Thandar (1989). 


Diagnosis.—Tentacles 10. Calcareous 
ring compact, short, tubular, with the radial 
and interradial plates fused for most of their 
length; posterior paired processes of the ra- 
dial plates of medium length, usually bro- 
ken into a few large pieces of calcite; rarely, 
processes unbroken. 

Type genus: Sclerodactyla Ayres. 


Pseudothyone Panning, 1949 


Diagnosis.—Tentacles 10. Calcareous 
ring undivided; radialia with medium length 
forked-tails, which are solid or in a few 
large pieces. Skin ossicles consist entirely 
of plates. 

Type species: Pseudothyone raphanus 
Diiben & Koren, 1846 


Pseudothyone levini new species 
Figs. 1—3 


Diagnosis.—Ten equal tentacles. Body 
cylindrical; length to 140 mm. Podia scat- 
tered but more numerous and regular in the 
ambulacra. Majority of skin ossicles thin, 
round smooth plates, with smooth wavy 
edges and a few small holes. Less common 
are round to oval plates with scalloped mar- 
gins and many large perforations. Tentacle 
ossicles range from small curved, lacy per- 
forated plates with scalloped margins to 
large, heavy curved rods. Radials of calcar- 
eous ring with short curved posterior pro- 
cesses; posterior edge of interradials con- 
cave; radials and interradials about three 
times longer than wide. 

Material examined.—Twelve specimens 
from nine localities. Material borrowed 
from: California Academy of Sciences 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(CASIZ), Canadian Museum of Nature 
(CMNI), Friday Harbor Laboratories 
(FHL), and the Royal British Columbia 
Museum (RBCM). Depths in meters. 

Holotype.-—RBCM 999-475-1, collected 
by Andy Lamb with aid of SCUBA, 13 
Nov 1999, length 14 cm, female. 

Type locality.—Point Defiance, Tacoma, 
Washington, USA, 47°18.4'N, 122°30.8'W, 
12 m, muddy sand. 

Paratypes.—RBCM 990-393-22, collect- 
ed by EW. Schueler, 4 Feb 1989, British 
Columbia, Queen Charlotte Islands, Gra- 
ham Island, Jungle Beach, 3 km south of 
Lawnhill, intertidal, 1 specimen, 3 cm long. 
RBCM 984-223-1, collected by D.B. Quay- 
le, 8 May 1961, British Columbia, Queen 
Charlotte Islands, Hecate Strait, Burnaby 
Narrows, 52°20.4'N, 131°20.4'W, intertidal, 
1 specimen, length 8.0 cm, male. RBCM 
980-343-20, collected by P. Lambert, 6 Jul 
1980, British Columbia, Kyuquot Sound, 
Rugged Point Light, 49°58.3'N, 127°15'W, 
<20 m on rock, | specimen, 11 cm long, 
male. RBCM 978-315-19, collected by P. 
Lambert, 26 Oct 1978, British Columbia, 
Gulf Islands, Prevost Passage, 48°43'N, 
123°20'W, 55—64 m in gravel, 1 specimen, 
12 cm long, male. FHL 819, collected by 
S. Van Neil, 13 Aug 1964, Washington, Or- 
cas Island, Potato Patch [local name], 
48°34.9'’N, 122°50.8’W, 40-55 m, 1 speci- 
men, length 9 cm, female. CASIZ 057397, 
collected by Mr. and Mrs. Oldroyd, Jul 
1917, Washington, San Juan Island, Friday 
Harbor, 48°32.7'N, 123°0.2'’W, est. 46-110 
m, | specimen, length 4.5 cm, male. RBCM 
996-193-1, collected by A. Lamb, 4 Jul 
1996, Washington, San Juan Island, Friday 
Harbor, 48°32.6'’N, 123°0.7'W, 12-18 m in 
soft mud, | specimen, length 9 cm, male. 

Other material.—RBCM 975-34-3, col- 
lected by John Fleury, 15 Aug 1962, British 
Columbia, Queen Charlotte Islands, Hous- 
ton Stewart Channel, 52°6.7'N, 131°8.2’W, 
30 m in gravel, 1 partial specimen, 1 cm 
posterior end. CMNI 1980-2097, collected 
by N.A. Powell, 17 Jul 1967, British Co- 
lumbia, Vancouver Island, Strait of Geor- 


VOLUME 114, NUMBER 3 


591 


Fig. 1. 
Tacoma, Washington, USA. 


gia, Nanoose Bay, 49°15.3’N, 124°8.0'W, 
19—24 m, in gravel, 2 partial specimens, an- 
terior end 3.5 cm and posterior end 2 cm 
long. CMNI 1980-2192, collected by N.A. 
Powell, 18 Aug 1967, Nanoose Bay, 
dredged parallel to the north end of the har- 
bor, 49°15.6'N, 124°10.0’W, 9-15 m in 
gravel, 3 partial specimens, 4 cm anterior 
end, two posterior ends 2.4 cm and 3.4 cm 
long. CMNI 1980-2107, collected by D. 
Popham, Aug 1967, British Columbia, Haro 
Strait, Moresby Island, 48°45.0’N, 
123°20.4'W, assumed to be a D.V. Ellis Sta- 


Holotype of Pseudothyone levini collected by Andy Lamb from a mud bottom near Point Defiance, 


tion, 70 m, 1 specimen, anterior piece 3 cm 
long. 

Description.—Body cylindrical, tapering 
gradually to posterior. Length of eight pre- 
served specimens, 3-14 cm (Holotype 9 
cm) with soft fleshy skin. In alcohol, skin 
yellowish white, flesh slightly darker. No 
obvious difference in color of ventral and 
dorsal surfaces. Tentacles white to dark 
brown. Living holotype beige with dark 
brown tentacles and purplish introvert (Fig. 
1). Ambulacral podia in five regular qua- 
druple rows and also thinly scattered be- 


592 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


rahe 
im 


Fig. 2. Top left: typical small-holed skin ossicles. Top right: supporting rods from podia. Mid left: large- 
holed skin ossicles. Mid right: plates and branched rods of the introvert. Bottom left: rods and plates from 
tentacles. Bottom right: calcareous ring, madreporite and polian vesicle. Upper scale = 100 pm. Lower right 
scale = 5 mm. 


VOLUME 114, NUMBER 3 


Fig. 3. Known distribution of Pseudothyone levini 
based on museum records. 


tween the ambulacra. Podia usually flush 
with the skin surface in preserved speci- 
mens. 

Ten dendritic tentacles; usually equal; en- 
circling the mouth. Eight specimens have 
ten equal tentacles; two have eight equal 
and two small ventral tentacles; and one has 
nine equal tentacles. Two small tentacles of 
One specimen white, eight larger tentacles 
brown. 

One polian vesicle in holotype, from zero 
to four in others. Usually one madreporite, 
but one specimen had a stone canal attached 
near calcareous ring and a second one pos- 
terior to that. Madreporite suspended in the 
dorsal mesentery (Fig. 2) but varying in po- 
sition between the midpoint of the calcar- 
eous ring and 15 mm posterior to the tips 
of the ring. Madreporic body ranging from 
globular and furrowed, as in holotype, to 
oblong and smooth. Mean length of mad- 
reporic body 1.9 + 0.6 mm, width 1.2 + 
0.4 mm. 

Radialia with short posterior processes; 
some are solid but others in small pieces. 
Other parts of calcareous ring variably solid 
or in pieces. Posterior edges of the interra- 
dials concave. Radials and interradials 
about three times longer than wide (Fig. 2). 


593 


Immediately posterior to each interradial, 
all specimens have a characteristic depres- 
sion or pit in the membrane (Fig. 2). 

Five retractor muscles attach to body 
wall at one third of body length from an- 
terior end. Holotype has wide, entire retrac- 
tors; in two specimens, all the retractors 
split longitudinally; in one specimen, one of 
the five retractors is split. Gonad two tufts 
of simple unbranched tubules, one on each 
side of dorsal mesentery. Each respiratory 
tree Y-shaped; dorsal branch often extend- 
ing as far as the tentacle bulb; ventral 
branch shorter. Base of trees join cloaca on 
each side of intestinal attachment. 

Generally two types of plates in skin. 
Commonly thin, round plates with smooth 
wavy margins and a few small holes; mean 
length 57 + 22 wm (n = 367); Holotype 65 
+ 11 (nm = 40); width 42 + 15 wm, Holo- 
type 50 + 8; mean number of holes 4 = 2, 
Holotype 4 + | with a maximum diameter 
of 6 = 4 my Holotype 10 = 3) (igs 2): 
Holes appear to get smaller with age, and 
some plates have none. Another type of 
plate, more common in smaller animals, 
round to oval and thick, with large holes 
and scalloped margins; mean length 80 + 
27 wm (7 = 147); width 56 + 17 wm; mean 
number of holes 6 + 3, with a maximum 
diameter of 13 + 5 pm (Fig. 2) Holotype 
has none of this type. Tentacular ossicles 
range from small, lacy perforated plates to 
large rods. Small perforated plates curved, 
with scalloped margins; vary from oval to 
triangular with mean length of 90 + 60 pm 
(n = 240), Holotype 84 + 20 (n = 40); 
wiglin 39) ae 22 ji, lnlolenjxs SS 22 IS, 
mean number of holes 9 + 7, Holotype 13 
+ 5, with a maximum diameter of 16 + 7 
wm, Holotype 15 + 5 (Fig. 2). Tentacular 
rods follow the circumference of the ten- 
tacle, thus vary in size, depending on lo- 
cation. Also, depending on orientation 
when viewed, they may appear narrow and 
curved with a few holes, or wide and heavy 
with many perforations (Fig. 2). Mean 
length of tentacular rods 212 + 123 wm (n 
= 280), Holotype 241 + 108 ( = 40). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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861 


‘roruuoqiey, 
pauodjnos “q 


VOLUME 114, NUMBER 3 


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596 


Introvert has small branched rods or ro- 
settes; mean length 65 + 23 pm (Fig. 2). 
No end plates found in podia; podia with 
branched rods similar to tentacles. Larger 
specimens have fewer ossicles in podia and 
skin; many have eroded edges. Deteriora- 
tion is likely natural rather than due to pre- 
servatives because tentacle ossicles in the 
same specimen not eroded. 

Etymology.—tThe species is named after 
Dr. Valery S. Levin, of the Kamchatka In- 
stitute of Fisheries and Oceanography, Pet- 
ropavlovsk, Russia in recognition of his nu- 
merous contributions to holothuroid sys- 
tematics and ecology. 

Distribution and bathymetric range.— 
Collections range from Graham Island, 
Queen Charlotte Islands, British Colum- 
bia (53°23’N, 131°55’W) to Point De- 
fiance, Tacoma, Puget Sound, Washington 
(47°18.4'N, 122°30.8’W) (Fig. 3). Depths 
range from intertidal to 70 meters, on rock 
or in mud or gravel. One record was be- 
tween 46 and 110 meters but the precise 
depth could not be confirmed. Most speci- 
mens occurred in less than 60 meters. Andy 
Lamb collected two specimens from mud 
with the aid of SCUBA. One specimen was 
collected in a subtidal exposed rocky hab- 
itat from a pocket of gravel and two spec- 
imens were collected from the intertidal 
zone. 


Discussion 


The family Sclerodactylidae is morpho- 
logically intermediate between the Phyllo- 
phoridae with complex calcareous rings and 
the Cucumariidae with simple rings (Paw- 
son 1966). Thandar (1989, 1990) states that 
several seemingly unrelated forms are in- 
cluded in the family, with some genera fit- 
ting equally well in the Phyllophoridae, 
which are characterized by long tubular cal- 
careous rings composed of a mosaic of mi- 
nute pieces. The family Sclerodactylidae 
currently contains three subfamilies: the 10- 
tentacled Sclerodactylinae and Sclerothyon- 
inae, largely Atlantic in distribution, and 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the Cladolabinae found typically in the 
Indo-West Pacific (Thandar 1989). Pawson 
& Fell (1965) state that having undivided 
processes is the key character of Sclerodac- 
tylidae that separates it from the Phyllo- 
phoridae; however, Thandar (1989) feels 
that this is a poor character because even 
within a single species of Sclerodactylidae 
the processes may be divided or undivided. 
He prefers to emphasize the short compact 
nature of the calcareous ring in this family 
(Thandar 1989, Fig. 1b). The calcareous 
ring of Pseudothyone levini is compact with 
short curved processes. While the processes 
in most specimens appear to be in small 
pieces, some are undivided. This variability 
in the posterior processes and the general 
character of the calcareous ring agree with 
the subfamily Sclerodactylinae as Thandar 
defined it. 

Table 1 compares the distribution and 
morphology of the new species with the 
seven known species of Pseudothyone, all 
of which have only plate ossicles in the 
body wall and generally inhabit mud in 
shallow water. Pseudothyone levini can be 
distinguished from other species of Pseu- 
dothyone by skin ossicles and the form of 
the calcareous ring. Pseudothyone levini of 
different sizes, and presumably different 
ages, show differences in the form of their 
skin ossicles as do specimens from rocky 
habitats versus mud. Specimens greater 
than 10 cm long had few of the large-holed 
plates found in small animals. This vari- 
ability in ossicles may represent growth 
stages, but a more complete size series of 
specimens is required before the relation- 
ship of size and ossicle form can be deter- 
mined. One specimen from a rocky sub- 
strate had larger small-holed skin ossicles 
than those from mud. Apart from the one 
rock-dwelling specimen the skin ossicles 
seemed to deteriorate in larger animals. 
This deterioration was unlikely a function 
of poor preservation as the tentacle ossicles 
in these specimens appeared normal. 

Pseudothyone levini appears to be most 
closely related to P. raphanus (Diiben & 


VOLUME 114, NUMBER 3 


Koren, 1846) from the North Atlantic. The 
skin ossicles of P. raphanus are a single 
type of perforated plate with large holes. 
Pseudothyone levini has a few large-holed 
perforated plates like the former but the 
dominant ossicles of mature specimens are 
smooth, rounded plates with small holes. 
The curved body of P. raphanus has an ex- 
panded anterior part and long slender cau- 
dal portion while P. levini is curved but 
only slightly tapered. Pseudothyone ra- 
phanus lives in soft substrates with its tail 
protruding and ingests the bottom material 
directly according to Mortensen (1977). On 
a night dive, Andy Lamb observed the ten- 
tacles of Pseudothyone levini extended 
above the mud surface presumably feeding 
on suspended particles. He collected one 
specimen and saw several other individuals 
in the same posture. On a second dive at 
the same location during daylight, none 
were seen. Several dredged specimens con- 
sist of only anterior and posterior portions, 
suggesting that the animal assumes a U- 
shaped position with anterior and posterior 
ends at the surface or protruding from the 
sediment. 

Four Pseudothyone species have calcar- 
eous rings with long posterior processes, 
three have medium length tails, and P. lev- 
ini has short curved tails. 

Unlike most specimens examined, two 
possessed small ventral tentacles. We spec- 
ulate that these may have been regenerating 
after an injury or in the case of the 3-cm 
specimen, may be a juvenile characteristic, 
but more specimens would be required to 
determine this. Tentacle sizes among the 
species of this genus vary (see Table 1). 
Either this character is variable in the genus 
or the genus is not well characterized and 
needs to be revised. We assume that varia- 
tions in the shape of the retractor muscles 
are artifacts that may have been caused by 
sudden contractions during preservation. 


Acknowledgments 


Thanks to Andy Lamb of the Vancouver 
Aquarium for his keen eyes in detecting un- 


597 


usual species and for collecting two speci- 
mens that helped us to complete the de- 
scription started a few years ago. The paper 
was improved by the constructive criticisms 
of Dr. Alex Kerr and one anonymous re- 
viewer. The authors thank the Director of 
the Royal B.C. Museum for research fund- 
ing and for supporting one of us (KO) dur- 
ing a University of Victoria Co-op work 
term in the fall of 1997 when the initial 
work on this new species was done. 


Literature Cited 


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cluding keys and descriptions to all continental 
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onomic atlas of the benthic fauna of the Santa 
Maria Basin and the Western Santa Barbara 
Channel, Santa Barbara Museum of Natural 
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Cherbonnier, G. 1958. Note sur Pseudothyone scul- 
ponea, nouvelle espece d’holothurie Dendro- 
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66. 

. 1969. Echinodermes recoltes par la Thalassa 
au large des cotes ouest de Bretagne et du Golfe 
de Gascogne (3-12 Aout 1967).—Bulletin du 
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serie 41:343-361. 

Clark, H. L. 1938. Echinoderms from Australia - an 
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Daniel, A., & B. P. Halder. 1974. Holothuroidea of the 
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ciation of India 16:412—436. 

Deichmann, E. 1930. The holothurians of the western 
part of the Atlantic Ocean.—Bulletin of the Mu- 
seum of Comparative Zoology (Harvard) 71: 
43-226. 

Diiben, M. W., von., & J. Koren. 1846. Ofversigt af 
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ska Vetenskaps-Akademiens Handlingar for 
1844:229-328. 

Froglia, C. 1975. Segnalazione di Pseudothyone scul- 
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ico di Storia Naturale Giacomo Doria 80:259— 
262. 

Koehler, R., & C. Vaney. 1910. Description d’holo- 
thuries nouvelles appartenant au Musee In- 
dien.—Records of the Indian Museum, Calcutta 
5:89—-104. 


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Lambert, P. 1985. Geographic variation of calcareous 
ossicles and the identification of three species 
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mata: Holothuroidea). Pp. 437-443 in B. E 
Keegan & B. D. S. O’Connor, eds., Echinoder- 
mata: Proceedings of the Fifth International 
Echinoderm Conference, Galway, A.A. Balke- 
ma, Rotterdam, 662 pp. 

. 1997. Sea cucumbers of British Columbia, 

Puget Sound and southeast Alaska. Royal BC 

Museum and University of BC Press, Vancou- 

ver, 192 pp. 


. 1998. Pentamera rigida and Pentamera pe- 
diparva, two new species of sea cucumber from 
the west coast of North America (Echinoder- 
mata: Holothuroidea).—Proceedings of the Bi- 
ological Society of Washington 111:535—550. 

Ludwig, H. 1886. Die von G. Chierchia auf der Fahrt 
der Kgl. Ital. Corvette ““Vettor Pisani” gesam- 
melten Holothurien.—Zoologiscen Jahrbtich- 
ern. Zeitschrift fiir Systematik, Geographie und 
Biologie der Thiere 1886:1—36. 

Mortensen, T. 1977. Handbook of the echinoderms of 
the British Isles. Dr. W. Backhuys, Uitgever, 
Rotterdam, 471 pp. 

Panning, A. 1949. Versuch einer Neuordnung der Fam- 
ilie Cucumariidae (Holothurioidea, Dendrochi- 
rota).—Zoologische Jahrbticher Abteilung fiir 


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Systematik, Oekologie und Geographie der Ti- 
ere 78:404—470. 

Pawson, D. L. 1966. Phylogeny and evolution of hol- 
othuroids. Pp. 641—646 in R. C. Moore, ed., 
Treatise on invertebrate paleontology, Part U, 
Echinodermata 3, vol. 2, Geological Society of 
America and University of Kansas Press, 
Lawrence, 1027 pp. 

. 1982. Holothuroidea. Pp. 813-818 in S. P. 

Parker, ed., Synopses and classification of living 

organisms, McGraw-Hill, Toronto, 1232 pp. 

, & H. B. Fell. 1965. A revised classification 
of the Dendrochirote Holothurians.—Breviora 
214:1-7. 

Sluiter, C. P 1910. Westindische Holothurien.—Zool- 
ogische Jahrbuecher Jena Suppl. 11:331—342. 

Stimpson, W. 1855. Descriptions of some new marine 
invertebrata. Philadelphia. 10 pp. 

Thandar, A. S. 1989. The sclerodactylid holothurians 
of southern Africa, with the erection of one new 
subfamily and two new genera (Echinodermata: 
Holothuroidea).—South African Journal of Zo- 
ology 24:290-304. 

. 1990. The phyllophorid holothurians of south- 
ern Africa with the erection of a new genus.— 
South African Journal of Zoology 25:207—223. 

Theel, H. 1886. Report on the Holothurioidea dredged 
by the HMS Challenger during the years 1873— 
1876 Part II. Voyage of the HMS Challenger- 
Zoology, vol. XIV, Part 39. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):599—604. 2001. 


A new leucosiid crab of the genus Nursia Leach, 1817 from Vietnam 
(Crustacea: Decapoda: Brachyura), with redescription of 
N. mimetica Nobili, 1906 


Hironori Komatsu and Masatsune Takeda 


(HK) Department of Biological Sciences, Graduate School of Science, The University of Tokyo, 
7-3-1 Hongo, Bunkyo-ku, Tokyo, 113-0033 Japan (corresponding address: Department of 
Zoology, National Science Museum, 3-23-1 Hyakunincho, Shinjuku-ku, Tokyo, 169-0073 Japan); 
(MT) Department of Zoology, National Science Museum, 3-23-1 Hyakunincho, Shinjuku-ku, 
Tokyo, 169-0073 Japan, and Department of Biological Sciences, Graduate School of Science, 
The University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo, 113-0033 Japan. 


Abstract.—A new species of leucosiid crab, Nursia guinotae, is described 
from Vietnam. It is similar to N. mimetica Nobili, 1906 from the Tuamotu 
Archipelago and N. alata Komatsu & Takeda, 1999 from the Ryukyu Islands, 
in the projecting epistome, the expanded epibranchial region, and the dorsal 
ridges, but is distinguished from them by the dentate epibranchial ridge and 
well-developed posterior lobes. Nursia mimetica, a species not reported since 
its original description, is redescribed based on the syntype specimen. 


Recently, we had the opportunity to ex- 
amine specimens of the genus Nursia 
Leach, 1817 deposited in the Muséum na- 
tional d’ Histore naturelle, Paris, through the 
courtesy of Prof. D. Guinot. Among this 
material, a small specimen which was col- 
lected in Vietnam and preliminarily identi- 
fied as N. plicata (Herbst, 1804) was found. 
Unfortunately, detailed locality of this spec- 
imen is unknown. This very small specimen 
has an expanded branchial region and a 
dentate epibranchial ridge, and actually rep- 
resents an undescribed species. Herein, we 
describe it as a new species, and redescribe 
an allied species, N. mimetica Nobili, 1906, 
from Rikitea, Tuamotu Archipelago, French 
Polynesia, to clarify the identity of both 
species, because the original and subse- 
quent descriptions of NV. mimetica by Nobili 
(1906, 1907) are very poor and insufficient. 

Measurements, given in millimeters 
(mm), are of the greatest carapace length 
(including the posterior lobe) and breadth, 
respectively. The descriptive terminology 
follows Ihle (1918), including the abbrevi- 
ations R and T in abdominal formulae 


which indicate Rest and Telson in German, 
respectively. The specimens remain depos- 
ited in the Muséum national d’ Histoire na- 
turelle, Paris (MNHN). 


Family Leucosiidae Samouelle, 1819 
Nursia guinotae, new species 
Figss 1.2 


Material examined.—Holotype, female, 
2.7 X 3.2 mm, Vietnam, coll. Gallardo, 
MNHN-B 9947. 

Description of holotype.—Carapace 
(Figs. 1, 2a, b) rhomboidal in general out- 
line, 1.3 times broader than long, uniformly 
covered with minute, flat granules, provided 
with postfrontal, median, and epibranchial 
ridges. Front well developed, 0.3 times as 
broad as carapace; margin slightly ridged, 
divided into 2 truncate lobes by shallow 
median notch; postfrontal ridges rising from 
near frontal-orbital angles, converging, 
meeting at midline. Orbit with 2 vestigial, 
longitudinal fissures on dorsal roof and V- 
shaped notch on infraorbital lobe. Ptery- 
gostomian margin forming general outline, 


600 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 
Scale equal 1 mm. 


with small angle near posterior end. Hepatic 
region faintly defined; margin inside gen- 
eral outline of carapace, angled near pos- 
terior end. Mesogastric median ridge gentle, 
narrow, rising from junction of postfrontal 
ridges. Gastric region with pair of tubercles 
and 1 median pearl-like granule slightly an- 
terior to tubercles. Cardiac region medially 
prominent, faintly separated from intestinal 
region by shallow transverse groove. Intes- 
tinal region strongly prominent, with me- 
dian ridge, culminating slightly anterior to 
posterior end. Epibranchial region strongly 
expanded, sloping anteriorly from ridge; 
margin thin, upturned, rounded, somewhat 
incurved near posterior end, forming right 
angle with metabranchial margin; epibran- 
chial ridge dentate, obliquely extending 
from near gastric tubercle to posterolateral 
margin, prominent at middle and distal end; 
prominences on epibranchial ridge, gastric 
tubercle, and gastric median granule almost 
coming into line. Metabranchial region 
deeply concave between epibranchial and 
intestinal ridges; margin weakly upturned, 
incurved at anterior 0.7, obtusely angled at 
posterior 0.3. Posterior margin trilobate; lat- 
eral lobes triangular with rounded tip, well 


Nursia guinotae, new species, holotype, ovigerous female, Vietnam, 2.7 X 3.2 mm (MNHN-B 9947). 


developed far beyond median lobe; median 
lobe small, triangular, with rounded tip, sit- 
uated slightly lower than lateral lobes. 

Third maxilliped (Fig. 2c, d) covered 
with minute granules; ischium 1.2 times as 
broad as exopod in maximum breadth; mer- 
us 1.2 times as long as ischium along me- 
sial margin; exopod arcuate and rimmed 
with short setae along lateral margin, with 
larger granules on distal part; internal me- 
sial ridge vestigial, with mesially directed, 
long setae. 

Cheliped (Fig. 2e) stout, entirely covered 
with granules of various sizes, sparsely 
frilled with short setae; merus subcylindri- 
cal, granules large and acute near inner and 
outer margins; carpus short, arcuate on out- 
er margin; palm medially swollen, some- 
what arcuate on inner margin, outer margin 
divided into 3 lobes, as long as movable 
finger along outer margin; both cutting edg- 
es of fingers with several minute, obtuse 
teeth, gaping and blunt at proximal 0.3. 

Ambulatory legs (Fig. 2f) slender, similar 
in shape, gradually decreasing in length 
from first to fourth, covered with minute 
granules except dactyli; meri subcylindri- 
cal, slightly longer than combined length of 


VOLUME 114, NUMBER 3 


carpi and propodi, with sparse soft setae; 
carpi slightly shorter than propodi; dactyli 
1.2 times as long as propodi, with vestigial, 
inconspicuous dactylo-propodal locks near 
proximal borders on dorsal surfaces (Fig. 
2g). 

Abdomen (Fig. 2h) entirely covered with 
minute granules on ventral surface; formula 
2+R+T; first segment completely con- 
cealed beneath carapace; second segment 
short, transversely zonal, bent proximally at 
median part; main fused section composed 
of third to sixth segments, ovoid, convex 
ventrally, divided into subregions by 2 lon- 
gitudinal and 3 transverse grooves, with ob- 
tuse triangular prominence near distal bor- 
der; telson tongue-shaped, directed dorsally 
in natural position. 

Etymology.—This species is dedicated to 
Prof. D. Guinot, who kindly gave the au- 
thors the opportunity to examine many 
valuable specimens deposited in the Musé- 
um national d’ Histoire naturelle, Paris. 

Color.—In preserved condition, body 
and appendages generally light brown; 
ridges, margins and posterior lobes of car- 
apace off white. 

Remarks.—This species is similar to N. 
alata Komatsu & Takeda, 1999 from the 
Ryukyu Islands, Japan, in the strongly ex- 
panded epibranchial region, the develop- 
ment of the frontal region, and the deeply 
concave metabranchial region. It is, how- 
ever, distinguished from WN. alata as fol- 
lows: the epibranchial ridges are dentate, 
whereas in N. alata the ridges are contin- 
uous; the mesogastric ridge is narrow and 
gentle, whereas in N. alata the ridge is 
broad, with a median shallow groove; the 
posterior lobes are more strongly expanded, 
whereas in N. alata the lobes are moderate- 
ly expanded. The new species also resem- 
bles N. mimetica Nobili, 1906, but is dis- 
tinguished from the latter by the dentate 
epibranchial ridge, the small projection of 
the pterygostomian margin, and the well- 
developed posterior lobes. 

Nursia guinotae, N. alata, and N. mime- 
tica can be distinguished from N. plicata 


601 


(Herbst, 1804) as follows: the hepatic and 
transverse ridges of the carapace are absent, 
whereas in N. plicata the ridges are present; 
the posterior margin of carapace is trilobate, 
whereas in WN. plicata the posterior margin 
is bilobed; the abdominal formula of female 
is 2+R+T, whereas in N. plicata the for- 
ame TS PZ ar Salar 1 


Nursia mimetica Nobili, 1906 
Fig. 3 


Nursia mimetica Nobili, 1906:261 (type lo- 
cality: Rikitea, Tuamotu Archipelago); 
1907: 380, pl. 1113); Komatsu & Takeda, 
IPOOe ine, WD), 


Material examined.—Syntype, 1 ovig. 
female, 2.7 < 3.7 mm, off Rikitea, Man- 
garéva I., Gambier Is., Tuamotu Archipel- 
ago, French Polynesia, 4—5 m, 1905, coll. 
G. Seurat, MNHN-B 17102. 

Redescription.—Carapace (Fig. 3a) 
transversely rhomboidal in general outline, 
1.4 times broader than long, closely cov- 
ered with minute granules, provided with 
postfrontal, median, and epibranchial ridg- 
es. Front moderately developed, 0.2 times 
as broad as carapace; margin slightly 
ridged, divided into 2 truncate lobes by 
small median notch; postfrontal ridges 
obliquely converging from frontal-orbital 
angles but not meeting together. Orbit with 
2 grooved, longitudinal fissures on dorsal 
roof; infraorbital lobe bearing V-shaped 
notch, with terminal sharrow pit. Pterygos- 
tomian margin forming general outline, al- 
most straight, roundly angled at posterior 
end. Hepatic region faintly defined; margin 
inside general outline of carapace, obtusely 
angled near posterior end. Mesogastric 
ridge broad, with longitudinal, shallow 
groove along midline, with some pairs of 
small pits on both sides of posterior end of 
ridge. Gastric region gently convex bilat- 
erally. Cardiac region weakly separated 
from intestinal region by faint transverse 
groove. Intestinal region gently swollen, 
with faint median ridge extending from car- 
diac region. Epibranchial region strongly 


602 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Lig, 2. 
a, carapace, dorsal view; b, same, frontal view; c, right third maxilliped, external view; d, same, internal view; 
e, right cheliped, dorsal view; f, right first ambulatory leg, dorsal view; g, dactylus of same, lateral view; h, 
abdomen, ventral view. Scales equal 1 mm (a—b, e-f, h), and 0.5 mm (c—d, g). 


expanded, sloping anteriorly from epibran- 
chial ridge; margin thin, weakly upturned, 
separated from pterygostomian margin by 
small notch, obtusely angled at posterior 
0.4, forming rounded, obtuse angle with 
metabranchial margin; epibranchial ridge 
weak, extending from near gastric convex- 
ity to metabranchial margin, interrupted 
medially. Metabranchial region shallowly 
concave between intestinal and epibranchial 
ridges, with 2 pairs of shallow pits on both 


Nursia guinotae, new species, holotype, ovigerous female, Vietnam, 2.7 X 3.2 mm (MNHN-B 9947): 


sides of cardiac and intestinal regions; mar- 
gin gently converging on anterior 0.3, in- 
wardly arcuate on posterior 0.7. Posterior 
margin trilobate, not well developed; laterel 
lobes situated lower than metabranchial re- 
gion; median lobe small, situated lower 
than lateral lobes. 

Third maxilliped (Fig. 3b, c) entirely 
covered with minute granules of various 
sizes; ischium longitudinally convex in lat- 
eral 0.7; merus 1.2 times longer than ischi- 


VOLUME 114, NUMBER 3 


603 


Fig. 3. 
mm (MNHN-B 17102): a, carapace, dorsal view; b, left third maxilliped, external view; c, right maxilliped, 
internal view; d, right cheliped, dorsal view; e, abdomen, ventral view. Scale equal 1 mm (a, d—e), and 0.5 mm 
(b—c). 


um along mesial margin, arcuate on lateral 
margin; exopod slightly convex along mid- 
line, with row of large granules, arcuate on 
lateral margin; internal mesial ridge vesti- 
gial, with several mesially directed, long se- 
tae. 

Right cheliped (Fig. 3d) moderate, 1.1 
times longer than carapace, covered with 
granules of various size; merus subcylindr- 
ical, slightly arcuate on outer margin; car- 
pus short, convex dorsally; palm swollen; 
movable finger 1.4 times as long as palm 
along outer margin; both cutting edges of 


Nursia mimetica Nobili, 1906, syntype, ovigerous female, Rikitea, Tuamotu Archipelago, 2.7 < 3.7 


fingers furnished with several triangular 
teeth on distal 0.4, blunt on proximal 0.6. 
Left cheliped missing. 

Ambulatory legs missing. 

Abdomen (Fig. 3e) entirely covered with 
minute granules; formula 2+R-+T; first seg- 
ment completely concealed beneath cara- 
pace; second segment short, transversely 
zonal, proximal border gently convex me- 
dially; main fused section composed of 
third to sixth segments, ovate, slightly con- 
vex ventrally, with 3 medially interruped 
transverse grooves and 2 short translucent 


604 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Differences between Nursia mimetica Nobili, 1906, N. alata Komatsu & Takeda, 1999, and WN. 


guinotae, new species. 


N. mimetica N. alata N. guinotae 
Mesogastric ridge broad with median groove broad with median narrow 
groove 
Epibranchial ridge oblique oblique oblique and dentate 
Posterior margin indistinctly trilobate, lower trilobate distinctly trilobate 


than metabranchial mar- 


gin 


Distribution Tuamotu Archipelago 


bands between grooves, with subtriangular 
prominence just near distal border, proximal 
border convex medially; telson short, 
tongue-shaped, with short setae around 
apex. 

Remarks.—This species is known only 
from the specimens used by Nobili (1906, 
1907), two females, from off Rikitea, Tua- 
motu Archipelago. Nobili noted that they 
mimick fragments of the calcareous alga, 
Halimeda opuntia. Tan & Richer de Forges 
(1993) also reported some mimetic crabs 
belonging to the family Leucosiidae. Al- 
though Nobili recorded two females in his 
descriptions, only one ovigerous female 
(syntype, MNHN-B 17102) is now depos- 
ited in MNHN. We inquired with the Museo 
Regionale de Scienze Naturali, Torino, for 
the remainder of the syntype specimens, but 
they were not found there (L. Levi, in litt.). 
The present specimen agrees well with No- 
bili’s descriptions, but is missing all its legs 
except the right cheliped. 

The comparison of N. mimetica, N. alata, 
and N. guinotae is summarized in Table 1. 


Acknowledgements 


We wish to express our cordial thanks to 
Prof. Daniéle Guinot, MNHN, for provid- 
ing the specimens. Our grateful thanks are 
due to Dr. Lisa Levi, Museo Regionale di 
Scienze Naturali, Torino, for her kind re- 
sponse to our inquiry; also to Drs. Tatsuo 
Katagiri and Noriko Uetani, Tokyo Metro- 


Ryukyu Islands Vietnam 


politan Institute for Neuroscience, for pro- 
viding HK with a stereoscopic microscope; 
and to Dr. Masayuki Osawa, National Sci- 
ence Museum, Tokyo, for improving the 
manuscript and general help. 


Literature Cited 


Herbst, J. E W. 1804. Versuch einer Naturgeschichte 
der Kraben und Krebse nebst einer systema- 
tischen Beschreibung ihrer vershiedenen Arten, 
3(4). Berlin, 1—49, pls. 59-62. 

Ihle, J. E. W. 1918. Die Decapoda Brachyura der Si- 
boga-Expedition. III]. Oxystomata: Calappidae, 
Leucosiidae, Raninidae.—Siboga-Expeditie 
39b:159-322. 

Komatsu, H., & M. Takeda. 1999. A new leucosiid 
crab of the genus Nursia from the Ryukyu Is- 
lands.—Bulletin of the National Science Mu- 
seum, Tokyo, Series A, 25:59—64. 

Leach, W. E. 1817. The Zoological Miscellany, Being 
Descriptions of New or Interesting Animals, 
vol. 3. London, 151 pp. + pls. 121-149. 

Nobili, G. 1906. Diagnoses préliminaires de Crustacés, 
Décapodes et Isopodes nouveaux recueillis par 
M. le Dr G. Seurat aux fles Touamotou.—Bul- 
letin du Museum National d’ Histoire Naturelle, 
Paris 12:256—270. 

. 1907. Ricerche sui Crostacei della Polinesia. 
Decapodi, Stomatopodi, Anisopodi e Isopodi.— 
Memorie della Reale Accademia delle Scienze 
di Torino, Classe di Scienze Fisiche Matemati- 
che e Naturali, Serie 2, 57:351—430, pls.1-3. 

Samouelle, G. 1819. The Entomologist’s useful Com- 
pendium, or An introduction to the knowledge 
of British insects. London, 469 pp. 

Tan, C. G. S., & B. Richer de Forges. 1993. On the 
systematics and ecology of two species of mi- 
metic crabs belonging to the familiy Leucosi- 
idae (Crustacea: Decapoda: Brachyura).—Raf- 
fles Bulletin of Zoology 41:119—132. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):605-610. 2001. 


First record of the portunid crab Arenaeus cribrarius (Lamarck, 


1818) (Crustacea: Brachyura: Portunidae) in marine waters 
of Argentina 


Marcelo A. Scelzo 


Departamento de Ciencias Marinas. Facultad de Ciencias Exactas y Naturales. Universidad 


Nacional de Mar del Plata/CONICET. Funes 3350, 7600 Mar del Plata, Argentina. E-mail: 
mascelzo @ mdp.edu.ar 


Abstract.—tThe first record of the portunid crab Arenaeus cribrarius (Lamarck, 
1818) in marine waters of Mar del Plata, Buenos Aires province, Argentina 
(38°S, 57°W) is documented. Juveniles and preadult specimens of the species 
were captured in waters at depths between 6 and 10 m on sand and mud-sand 
bottom, 10°C of temperature and 34%oc of salinity. This report extends the 
southern limit of distribution of A.cribrarius by more than 500 km. This species 
was previously known in the southwestern Atlantic from Massachusetts to Uru- 
guay. 

Resumen.—E] trabajo documenta el primer hallazgo del cangrejo portunido 
Arenaeus cribrarius (Lamarck, 1818) en aguas marinas frente a Mar del Plata, 
provincia de Buenos Aires, Argentina (38°S, 57°W). Los ejemplares juveniles 
y preadultos de la especie fueron capturados a profundidades entre 6 y 10 m 
sobre fondo arenoso y areno-fangoso, en aguas con temperaturas de 10°C y 
34%o de salinidad. Con esta menci6n se extiende la distribucion de A.cribrarius 
mas de 500 km. La especie es conocida previamente en el Atlantico Sudoc- 


cidental desde Massachusetts hasta Uruguay. 


In June 2000, Mr. José L. Ungarelli, a 
local fisherman captured three specimens 
of what appeared to him as a strange crab 
for the marine waters near Mar del Plata 
(38°S, 57°W), Buenos Aires province, Ar- 
gentina. The specimens were identified as 
Arenaeus cribrarius (Lamarck, 1818) 
(Fig. 1) a portunid crab known to range 
broadly from shallow waters of Vineyard 
Sound, Massachusetts, U.S.A., throughout 
the Caribbean Sea, to Brazil and Uruguay 
(Williams 1965, 1984; Juanicéo 1978; 
Melo 1996, 1998, 1999). The type locality 
is Brazil. 

This coastal species found in the Virgi- 
nian to Argentinean zoogeographical prov- 
inces in the southwestern Atlantic (Boschi 
2000a, 2000b) had not been previously cap- 
tured south of Uruguay. The present report 


in the Mar del Plata region (38°S), extends 
the distribution of A. cribrarius by more 
than 500 km from the previous southern 
limit in the southwestern Atlantic Ocean 
(Juanico 1978). 

The specimens were captured using a 
beam trawl at depths of 6 tol0 m over sand 
and mud-sand bottom. Water was 10°C in 
temperature, and 34%o in salinity. 

Specimens were measured for carapace 
width (CW) excluding lateral spines, and in 
carapace length (CL), from basal region be- 
tween the frontal teeth to the postero-me- 
dial border of carapace (Pinheiro & Fran- 
sozo 1993, 1998). Measurements are in mil- 
limiters, and were obtained with a caliper 
to the nearest 0.1 mm accuracy. Specimens 
were weighed (W) in a digital scale (0.001 
grams sensibility). 


606 


Family Portunidae Rafinesque, 1815 
Arenaeus cribrarius (Lamarck, 1818) 
Fis. | 


Portunus cribrarius.—Lamarck. 1818:259. 

Arenaeus cribrarius.—Williams 1965:173, 
fig. 153.—1984:362-363, fig. 292. 
Melo 1998:476.—1999:450, fig. 37. 


Material examined.—3 juveniles females 
CW 35.5, 36.5 and 40.2 mm, CL 20.2, 21.5 
and 24.0 mm, W 5.8, 6.59 and 8.82 g. Two 
small specimens were deposited in the col- 
lections of the Departamento de Ciencias 
Marinas, Universidad Nacional de Mar del 
Plata, and the larger specimen in the Museo 
Argentino de Ciencias Naturales ““Bernar- 
dino Rivadavia’’, Buenos Aires, Argentina 
(Collection number: MACN 34616). 

Remarks.—The specimens showed white 
spots characteristic of the species, and mor- 
phologically agree with the characters pro- 
vided by Williams (1984). The small crabs 
still showed a white stripe on the carapace, 
indicative of a juvenile stage. All three 
specimens are juvenile females, with the 
pleon plate still adhered to the sternum and 
they have a triangular shaped abdomen 
characteristic of juvenile stages as indicated 
by Pinheiro & Fransozo (1998). 


Discussion 


In recent years several new records of 
crustaceans from Argentinean marine wa- 
ters have been reported in papers and/or 
meetings. Lini et al. (1995) documented the 
first record of the isopod Joeropsis dubia in 
Mar del Plata; Spivak & Bas (1999) found 
the grapsid crab Planes marinus Rathbun, 
1914 in Mar Chiquita, 80 km north of Mar 
del Plata, together with other crustaceans 
such as the cirriped Lepas anatifera L., 
1758 and the amphipod Caprella andreae 
Mayer, 1890, all representing first records 
for the southwestern Atlantic Ocean. 

Several hypotheses have been proposed 
for the presence of these species, including 
ships fouling and ballast water, drifting 
plankton larvae, and surface current trans- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


port associated with floating objects. Ac- 
cording to specialists of fishes (L6pez 
1964), phytoplankton (Balech 1964), crus- 
taceans (Boschi et al. 1992, 2000a, 2000b) 
the Argentinean biogeographic province is 
the area between 23°S (Rio de Janeiro, Bra- 
zil) to 43°S (Rawson, Chubut). According 
to Lépez (1964) it is possible to distinguish 
two districts: southbrazilian and bona- 
erense, with a limit around 34°S, but that 
limit can be modified according movements 
of water masses, specially near the coasts, 
reaching near San Jorge Gulf, Argentinean 
Patagonia during summer months. Low sa- 
linity waters of Rio de la Plata are consid- 
ered as a natural barrier for the dispersion 
of marine fauna. As it was stated by Lucas 
et al. (1999:105). . .“‘the presence of winds 
that force onshore flow to the south seem 
to be favorable for the southward extension 
of low salinity water from Rio de la Plata 
and are most likely to occur in spring-sum- 
mer. Anomalously high discharge from the 
Rio de la Plata associated with ENSO (El 
Nifio south oscillation) conditions in spring 
of 1997 and summer of 1998 showed a cor- 
relation with a coastal low salinity intrusion 
as far south as Miramar’’... more than 40 
km south of Mar del Plata. Eventually, 
warm waters south to the bonaerense litto- 
ral would be associated to a intrusions of 
the Brazil Current (Balech 1965, 1986), wa- 
ter circular movements or eddies (Piola & 
Rivas 1997), or the so called Argentinean 
flux (waters of subtropical origin) according 
to Severov (1990). Other hypotheses pos- 
tulate that the presence of marine biota of 
southbrazilian origin in bonaerense coastal 
region will be caused by the diminishing 
caudal of fresh-water from Rio de la Plata, 
at it happened during summer and fall of 
the year 2000 (Mianzan et al. 2000). 

The number of brachyuran crabs living 
in littoral waters of Argentina reach 39 
species (Boschi 1964; Boschi et al. 1992; 
Spivak & Bas 1999), of which three are 
portunids: Ovalipes trimaculatus (de 
Haan, 1833), Coenophthalmus tridentatus 


VOLUME 114, NUMBER 3 607 


TTT 


Fig. 1. Arenaeus cribrarius juvenile: A) dorsal view, B) ventral view. Scale in millimeters. 


608 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


50 == a aan ee 0k 


08 


Mar del 
Plata_ bs 


60° 


140 120° “400 


Biss 2? 


Geographical distribution of Arenaeus cribrarius in the western Atlantic Ocean. 
record. 


* Indicates new 


VOLUME 114, NUMBER 3 


A.Milne Edwards, 1879 and Callinectes 
sapidus Rathbun, 1896. The presence of A. 
cribrarius adds another species to this re- 
gion. Arenaeus cribrarius lives in shallow 
waters along ocean beaches from the water 
line to 68 m (Williams 1984; Melo 1996, 
1998, 1999). Ovigerous females of this spe- 
cies are present during September in Ven- 
ezuela and Brazil (Williams 1984) and dur- 
ing all months in Ubatuba region (Pinheiro 
& Fransozo 1998). The presence of this 
species near Mar del Plata can be explained 
based on larval transport by plankton drift- 
ing from coastal subtropical waters during 
spring-summer, crossing Rio de la Plata af- 
ter metamorphoses, and then growing to 
CW of 35.5 to 40.2 mm; or larvae intro- 
duced by ballast water of ships, as men- 
tioned for other portunidae species in other 
localities of the western Atlantic and the 
Caribbean Sea (Lemaitre 1995; Tavares & 
Mendonga 1996; Mantelatto & Dias 1999). 
The specimens reported herein represent ju- 
venile stages, and thus it appears that the 
adult population is not yet established. The 
occasional finding of juveniles in marine 
waters of Mar del Plata, Argentina as well 
as in Uruguay by Juanico (1978) will be 
considered as expatriation areas in the dis- 
tribution of the species in the southern At- 
lantic ocean (Fig. 2). 


Acknowledgments 


I thank Dr. R. Lemaitre, NMNH, USA, 
for manuscript criticisms and English cor- 
rections. Author also thank to Dr. E. E. Bos- 
chi, INIDEP, Argentina and other anony- 
mous reviewers to their valuable criticisms 
to the manuscript. Special thanks to Mr. 
José Luis Ungarelli for obtaining the crabs 
and to Eng. J. Martinez Arca for his assis- 
tantship. Study granted by Projects: Univ- 
ersidad Nacional de Mar del Plata E 207/ 
2000, Consejo Nacional de Investigaciones 
Cientificas y Tecnologicas CONICET PIP 
4346/97 and Antorchas 13817—7/2000. Sci- 
entific Contribution Departamento de Cien- 
cias Marinas, UNMDP N° 105. 


609 


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Lini, R. M., A. M. Roux, & M. A. Scelzo. 1995. Sobre 
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):611—617. 2001. 


First record of larvae of the rare mud shrimp Naushonia Kingsley 
(Crustacea: Decapoda: Laomediidae) from Asian waters 


Kooichi Konishi 


National Research Institute of Aquaculture, Nansei, Mie 516-0193, Japan 


Abstract.—Larvae of an undetermined species of the laomediid genus Nau- 
shonia Kingsley are reported from plankton samples collected in Gokasho Bay, 
Pacific coast of Japan. The presence of stage 1 zoeas suggests that adult shrimps 
of Naushonia are distributed in the neighboring district. This is the first record 


of Naushonia from Asian waters. 


The rare laomediid genus Naushonia 
Kingsley, 1897 consists of seven species, 
none of which are known from Asian wa- 
ters (Martin & Abele 1982, Berggren 1992, 
Alvarez et al. 2000). There is scant larval 
information on these rare mud shrimps to 
date, although all known species of Nau- 
shonia occur in very shallow benthic wa- 
ters. Since Thompson’s (1903) work, plank- 
tonic larval stages of Naushonia species 
have been described for N. crangonoides 
Kingsley, 1897, N. portoricensis (Rathbun, 
1901) and two undetermined species (Gur- 
ney & Lebour 1939, Dakin & Colefax 
1940, Kurian 1956, Goy & Provenzano 
1978). 

During plankton collections made at Go- 
kasho Bay, Japan, in the summer of 2000, 
zoea | larvae belonging to an undetermined 
species of Naushonia were found. These 
zoeas are described in this study, represent- 
ing the first record of the genus in Asian 
waters. 


Materials and Methods 


The zoea 1 larvae were taken in a 300 
zm mesh and 46 cm diameter tow-net, at 
depths of 5-10 m, at Gokasho Bay, Mie 
Prefecture, Japan, 34°20'’N, 136°42’E, on 
27 Jul, 2 and 10 Aug 2000. Water temper- 
ature and salinity of the collection sites 
ranged from 25 to 27°C and 33.5 to 33.7%o 
respectively. 


Living specimens were observed and 
sorted under a Nikon SMZ-10 stereomicro- 
scope. After fixation in 3% buffered for- 
malin, the appendages were dissected with 
fine insect pins or sharpened tungsten nee- 
dles under the stereomicroscope, and 
mounted on a silicon-coated glass slide. 
Drawings and measurements were made 
with a drawing tube attached to an Olympus 
BH-2 microscope. A color image of a living 
specimen was captured with an Olympus 
HC-300/OL digital camera connected to the 
microscope. All illustrations were made 
with Illustrator 5.5J (Adobe Systems Inc.) 
on a Mactintosh® OS (Apple Co. Ltd.). 
Carapace length (CL) was measured from 
the tip of the rostral spine to the medial 
posterior border of the carapace. 

Voucher specimens have been deposited 
in the Natural History Museum and Insti- 
tute, Chiba, Japan, under accession num- 
bers CBM-ZC 5573-5575. 


Description of zoea 1 


Size: CL = 0.73 + 0.036 mm (range 
0.70—0.77 mm, 3 specimens). 

Color (Fig. 1): Small red and large yel- 
lowish chromatophores dispersed on anten- 
nule, and mainly ventral side of carapace 
and abdomen. 

Carapace (Fig. 2A, B): Rostral spine 
short, slender, upturned distally. Posterolat- 
eral border rounded. Eyes sessile. 


612 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 
Scale = 0.1 mm. 


Antennule (Fig. 3A): Uniramous, about 
half-length of CL, unsegmented with 5 
aesthetascs terminally, and long plumose 
seta subterminally. 

Antenna (Fig. 3B): Biramous. Protopod 
with simple spine at base of exopod. Endo- 
pod with 3 long plumose setae distally. Ex- 
opod (=antennal scale) flat, elongated, oval, 
with 10 plumose setae on inner margin. 

Mandibles (Fig. 3C): Asymmetrical. Left 
with incisor process large, sickle-shaped, 
with 2 subterminal inner spines; right small, 
conical, with subterminal inner spine. Right 
and left molar processes almost equal in 
size, but different in dentation. Paragnath of 
left side (Fig. 3C’) slender sickle-shape as 
in left mandible. 

Maxillule (Fig. 3D): Coxal endite with 4 
simple setae and subterminal small seta. 
Basial endite with 2 large teeth and 2 sim- 
ple setae. Endopod unsegmented with 3 ter- 
minal setae. 

Maxilla (Fig. 3E): Coxal endite bilobed, 
proximal lobe small, with 1 long and 1 
short plumose setae, and distal lobe with 
simple 3 setae. Basial endite bilobed with 
5+4 simple setae, respectively. Scaphog- 
nathite without proximal lobe, and with 5 
plumose setae on margin. 


Photograph of a living specimen of zoea 1 of Naushonia sp. Note chromatophores on ventral side. 


Maxilliped | (Fig. 3F): Biramous. Coxa 
without setae. Basis with 1+2+3 inner se- 
tae. Endopod 4-segmented with 2,1,2,5+1 d 
= dorsal seta) seta. Exopod with 4 natatory 
plumose setae terminally, symmetrically ar- 
rangement in 2 tiers (Fig. 3F’). 

Maxilliped 2 (Fig. 3G): Biramous. Coxa 
and basis with no setae. Endopod 4-seg- 
mented with 0,0,2,4+I setae. Exopod as in 
maxilliped 1. 

Maxilliped 3 (Fig. 3H): Uniramous, 3- 
segmented, without setae. 

Pereiopods: Rudimentary. 

Abdomen (Fig. 2A): Smooth, 5 somites 
plus forked triangular telson. Abdominal 
somite | with short rod-like pleural projec- 
tion (Fig. 2A’) and somite 2-5 with anteri- 
orly procurved pleural hooks. 

Telson (Fig. 2C, C’): Triangular, with 
deep notch and 7 pairs of posterior pro- 
cesses; outermost process stout, unarticulat- 
ed spine; second process thin plumose seta 
(=anomuran hair) on ventral side of telson- 
al fork (Fig. 2C’); third through seventh 
processes large articulated plumose setae. 


Discussion 


The infraorder Thalassinidea consists of 
11 families (Poore 1994) and larval stages 


VOLUME 114, NUMBER 3 613 


i 


EET Tf 


Fig. 2. Naushonia sp., zoea 1. A, whole animal, lateral view; A’, enlarged posterior part of abdominal somite 
1; B, carapace, dorsal view; C, telson, dorsal view; C’, enlarged right telsonal fork, ventral view, showing 
‘anomuran hair’ (arrow). 


614 PROCEEDINGS OF 


oe 


—— 


THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Naushonia sp., zoea 1. A, antennule; B, antenna; C, mandibles; C’, left paragnath lobe; D, maxillule; 
E, maxilla; EK maxilliped 1; F’, tip of exopod of maxilliped 1; G, maxilliped 2; H, maxilliped 3. 


have been described for species in six fam- 
ilies. Laomediid zoeal larvae differ from 
those of other decapod shrimps by the con- 
siderably asymmetrical mandibles and pro- 
curved pleural hooks on the abdominal so- 
mites (see Konishi 1989). A recent study of 


Thalassina anomala (Herbst, 1804) of the 
Thalassinidae Latreille, 1831, has revealed 
that its zoea also has a sickle-shaped asym- 
metrical pair of mandibles as in the La- 
omediidae Borradaile, 1903 (Uchino 1993). 
Zoeas of T. anomala, however, have a 


VOLUME 114, NUMBER 3 


615 


Jaxea 


lLaomedia 


Naushonia 


Axianassa 


Fig. 4. Diagrammatic representation of zoeal cephalothoraxes in four laomediid genera showing relative 
position of mandible (shaded). Vertical dotted line indicates midpoint of carapace length. (Based on Gurney 


1924; Ngoc-Ho 1981; Fukuda 1982). 


unique larval character: 1.e., the extremely 
elongated endopod of maxilliped 2. Thus, 
the laomediid zoeas can be easily separated 
from the Thalassinidae despite similarities 
in the mandibles. 

The Laomediidae includes 5 genera as 
follows (Ngoc-Ho 1981): Laomedia De 
Haan, 1841, MNaushonia, Jaxea Nardo, 


1847, Axianassa Schmitt, 1924, and Laur- 
etiella Le Loeuff & Intes, 1974. Larval 
stages are known for representative species 
in all of them except Lauretiella (Thomp- 
son 1903, Caroli 1924, Gurney 1924, Gur- 
ney & Lebour 1939, Dakin & Colefax 
1940, Kurian 1956, Sakai & Miyake 1964, 
Goy & Provenzano 1978, Ngoc-Ho 1981, 


616 


Fukuda 1982, Rodrigues & Shimizu 1992). 
Their zoeas are clearly distinguishable be- 
tween genera. The mandibles of Naushonia 
and Axianassa are located anterior to the 
mid point of the carapace as in typical deca- 
pod zoeas, while those of Jaxea and La- 
omedia are posterior (Fig. 4). The rostrum 
of Naushonia is short and upturned whereas 
it is long and straight in Axianassa. In ad- 
dition, the abdomen of Axianassa zoea bear 
a pair of posterolateral spines on somite 5 
instead of procurved pleural hooks on so- 
mite 2-5 as found in Jaxea, Naushonia and 
Laomedia. Therefore, zoeas of Naushonia 
species are easily identified. 

The morphology of the herein reported 
Japanese zoeas is different from those of WN. 
crangonoides and N. portoricensis de- 
scribed by Goy & Provenzano 1978 and 
Gurney & Lebour 1939, in antennule, max- 
ilule and pleural projection on abdominal 
somite 1. The antennule, has two subter- 
minal plumose seta in N. crangonoides, 
whereas only one in the present zoea. The 
coxal endite of maxillule bears a small 
proximal lobe in the present zoea, whereas 
there is no lobe in the zoea of others. The 
present zoeal specimens, therefore, seems 
to belong to a species different from those 
for which larvae are known, or perhaps to 
an unknown species of the genus. 

Until now, the Laomediidae was repre- 
sented in Japan by only one genus Laome- 
dia (Miyake, 1998). The present zoeas, 
however, are clearly assignable to Naushon- 
ia species. According to findings for rearing 
of plankton-caught zoeas of N. crangono- 
ides by Goy & Provenzano (1978), zoea 1 
molt to the next instar within 5.24 days in 
average when they were incubated at 25°C. 
This strongly suggests that adult shrimps of 
Naushonia must be distributed in the neigh- 
boring district of Gokasho Bay in Japan or 
very nearby waters. 

There may be two reasons why Nau- 
shonia larvae have not been reported pre- 
viously from Japan and adjacent waters. 
First, they may be of very rare species 
(Berggren 1992) in Japan and adjacent 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Asian waters. Second, Naushonia does not 
have native representative, and the present 
zoeas are from recent introduction of an ex- 
otic species, as has occurred with the Med- 
iterranean portunid crab in Tokyo Bay, Ja- 
pan (Sakai 1986). A careful faunal survey, 
and further larval studies of the coast of 
Japan, may be needed to answer this ques- 
tion. 


Acknowledgements 


The author thanks Dr. R. Lemaitre (Smi- 
thonian [nstitution) for critical reading of 
the manuscript; and Dr. H. Yokoyama, Y. 
Ishihi and S. Yamamoto (National Research 
Institute of Aquaculture), for their help in 
collecting the plankton. My cordial thanks 
are due to Dr. T. Komai (Natural History 
Museum and Institute, Chiba) for valuable 
information about systematics of adult 
shrimp. 


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VOLUME 114, NUMBER 3 


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40:237-273. 

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):618-623. 2001. 


A new cavernicolous species of freshwater crab (Crustacea: 
Brachyura: Potamidae) from Pulau Tioman, peninsular Malaysia 


Darren C. J. Yeo 


Raffles Museum of Biodiversity Research, Department of Biological Sciences, 
National University of Singapore, 10 Kent Ridge Crescent, Singapore 119260. 


Abstract.—A new freshwater potamid crab species, Johora gua, is described 
from Pulau Tioman, an island off the east coast of Peninsular Malaysia. It differs 
from most of its congeners by the relatively longer and slenderer ambulatory 
legs as well as various other characters in the carapace and male first pleopod. 
This is the fifth species of Johora Bott to be described from Pulau Tioman, 
and the first cavernicolous crab to be reported from Peninsular Malaysia. 


Specimens of an unusual freshwater crab 
recently collected from a cave on Pulau 
Tioman, an island off the east coast of Pen- 
insular Malaysia, proved to be a new po- 
tamid species of the genus Johora Bott, 
1966. This new species differs from most 
of its congeners by the relatively longer and 
slenderer ambulatory legs as well as various 
other characters of the carapace and male 
first pleopod. The genus Johora Bott, 1966 
[type species: Potamon (Potamon) johor- 
ense Roux, 1936], is endemic to the Malay 
Peninsula, occurring northwards from Sin- 
gapore into the southern half of Peninsular 
Malaysia along the central highlands. Spe- 
cies of this genus are replaced by those of 
Stoliczia Bott, 1966, in the northern half 
through to southern Thailand (Ng 1988). A 
total of 13 species have been placed in Jo- 
hora, with Pulau Tioman alone accounting 
for four species, all island endemics, viz., 
J. counsilmani (Ng, 1985), J. grallator Ng, 
1988, J. punicea (Ng, 1985), and J. tiom- 
anensis (Ng & Tan, 1984) (see Ng 1988, 
1990; Ng & Takeda 1992; Yeo et al. 1999). 
In the present study, the fifth Johora species 
known from Pulau Tioman is described, 
bringing the total number in the genus to 
14. 

Ng (1988: 142) earlier referred to the 
*,..existence of an unknown species of 
white crab in Malayan caves...” but this 


“6 


has yet to be discovered. The new species 
of Johora herein described is the first true 
cavernicolous freshwater crab to be report- 
ed from Peninsular Malaysia. The cave- 
dwelling habits of this species are, to some 
extent, reflected in slight troglomorphic fea- 
tures although these are less significant 
When compared with other known caver- 
nicolous crabs. 

The following abbreviations are used: 
G1, male first pleopod; G2, male second 
pleopod; cw, carapace width; and asl, above 
sea level. Measurements are given in mil- 
limeters (mm) as carapace width X length. 
Terminology used essentially follows Ng 
(1988). Malay words used in the text are 
‘Pulau’, island; “Gua’, cave; and ‘Gunung’, 
mountain. Specimens remain deposited in 
the Zoological Reference Collection, Raf- 
fles Museum of Biodiversity Research, De- 
partment of Biological Sciences, National 
University of Singapore (ZRC). As com- 
parative material, the types and supplemen- 
tary specimens of the other four Johora 
species from Pulau Tioman in the ZRC 
were examined. A detailed listing of this 
material can be found in Yeo et al. (1999). 


Johora gua, new species 
Figs. [A—C, E—H, 2A—G 


Material examined.—Holotype: male 
11.4 X 8.5 mm (ZRC 2000.2236), Gua 


VOLUME 114, NUMBER 3 


Tengkuk Ayer, route to Gunung Kajang 
from Juara, Pulau Tioman, ca. 900 m asl., 
Peninsular Malaysia, coll. H. H. Tan & T. 
M. Leong, 1 Sep 2000. Paratype: male 9.5 
xX 7.2 mm (ZRC 2000.2237), same data as 
holotype. 

Diagnosis.—Carapace distinctly broader 
than long, not elevated; dorsal surface flat, 
glabrous. Epigastric cristae distinctly ru- 
gose; postorbital cristae not sharp, rugose. 
Frontal region smooth; antennular fossae 
broadly subrectangular; corneas slightly re- 
duced, about 0.4 times length of eyestalks. 
External orbital angle low, broadly trian- 
gular, outer or lateral margin about 4 times 
longer than inner or medial margin; epi- 
branchial tooth weak, triangular; anterolat- 
eral margin gently convex, very weakly 
cristate; branchial region indistinctly ru- 
gose; metabranchial region with distinct 
oblique striae. Epistome with weak, broadly 
triangular median tooth on posterior mar- 
gin. Third maxilliped exopod with well-de- 
veloped flagellum, longer than merus width. 
Cheliped with carpus armed with robust 
subdistal spine on inner margin. Ambula- 
tory legs sparsely setose, long, slender; dac- 
tyli elongated, slender, merus unarmed. Su- 
ture between anterior thoracic sternites 2 
and 3 faint; thoracic sternite 8 completely 
separated by longitudinal median line, lack- 
ing transverse ridge on midline; abdominal 
cavity reaching imaginary line joining an- 
terior edge of cheliped bases. Male abdo- 
men triangular. G1 terminal segment about 
0.4 times length of subterminal segment, 
very slender, subcylindrical, curved out- 
wards, lacking dorsal flap; subterminal seg- 
ment distinctly broader in proximal half, 
not neck-like distally, without shelf or cleft 
on upper part of outer margin. G2 distal 
segment distinctly longer than half of basal 
segment. 

Color.—Live specimens are light orange 
in overall coloration. 

Etymology.—tThe specific name, “‘gua’’, 
is Malay for cave, alluding to its cavernic- 
olous habitat. Used as a noun in apposition. 

Ecological notes.—The specimens of Jo- 


619 


hora gua, new species, were obtained sev- 
eral meters inside a small granite cave, not 
higher than 1 m in height, with a shallow 
underground stream, about 2-15 cm deep, 
flowing slowly over sandy and rocky sub- 
stratum. The water pH was 7.8. No surface 
(epigeal) streams were found in the vicinity 
(T. M. Leong & H. H. Tan, pers. comm.; 
Ng et al. 1999: 171). Organic input into this 
system include swiftlet and porcupine drop- 
pings. Tadpoles of a species of megophyrid 
frog, Leptolalax gracilis (Giinther), and a 
species of fish, the blind cave loach, Sun- 
doreonectes tiomanensis Kottelat, were 
found syntopically, along with the fully 
aquatic freshwater crab species J. tioma- 
nensis (I. M. Leong & H. H. Tan, pers. 
comm.). Despite its presence in the cave, 
and the absence of surface streams nearby, 
J. tiomanensis 1s still regarded as an epigeal 
species, as it is commonly found in surface 
streams at lower altitudes throughout the 
southern half of the island (see Yeo et al. 
1999). 

Remarks.—Johora gua, new species, is 
immediately separated from all its conge- 
ners except J. grallator Ng, 1988, by its 
much longer and slenderer ambulatory legs. 
Johora gua could be mistaken for a juvenile 
of J. grallator, as they are superficially sim- 
ilar in the long, slender legs as well as light 
orange live coloration (see Yeo et al. 1999). 
However, specimens of J. gua are much 
smaller, similar in size to the holotype (11.4 
xX 8.5 mm, ZRC 2000.2236), which clearly 
possesses well-developed adult Gls. In 
contrast, two similar sized male specimens 
of J. grallator (largest 11.7 X 8.9 mm, ZRC 
1996.1730) have undeveloped Gls and are 
clearly juvenile. The ambulatory legs of J. 
gua are relatively stouter than those of J. 
grallator, as reflected in the proportions of 
the merus of the second leg (4.3—4.4 vs. 
5.3—5.7 times longer than broad); and mer- 
us of the fourth leg (3.4—3.6 vs. 4.4—4.7 
times longer than broad) (Fig. 2A, B; Ng 
1988: fig. 23A). Furthermore, J. gua differs 
from J. grallator in the following charac- 
ters: postorbital cristae relatively more ru- 


620 


gose and less sharp (vs. postorbital cristae 
distinctly smoother and sharper); corneas 
reduced, shorter than half the length of the 
eyestalks (vs. normal-sized corneas, about 
half the length of the eyestalks); antennular 
fossae distinctly broader than the epistome 
(vs. antennular fossae as broad as epi- 
stome); and epistome posterior margin with 
poorly developed median tooth (vs. epi- 
stome posterior margin with well-devel- 
oped median tooth) (Fig. 1A, C; Ng 1988: 
ie, 22vA, IBS VEO Gt ail, WILLE ily, Sys\; 383), 
The shape of the G1 of J. grallator is un- 
known, as no mature males have been col- 
lected (see Ng 1988; Yeo et al. 1999). How- 
ever, the external morphological differences 
discussed are sufficient to show that the two 
species are clearly not conspecific. Most of 
these external differences are also applica- 
ble in distinguishing J. gua from J. tioma- 
nensis (Ng & Tan, 1984), and J. counsil- 
mani (Ng, 1985), two other large, fully 
aquatic Johora species that are endemic to 
Pulau Tioman (see Ng 1988: figs. 20A, B, 
21A, B). In addition, J. gua can be further 
separated from these species by its almost 
smooth sub-branchial, sub-hepatic and pter- 
ygostomial regions (vs. sub-branchial, sub- 
hepatic and pterygostomial regions distinct- 
ly rugose); and by its G1 terminal segment 
always lacking a dorsal flap (vs. G1 ter- 
minal segment with dorsal flap); and Gl 
subterminal segment lacking a distal cleft 
on the outer margin (vs. G1 subterminal 
segment with a distinct subrectangular dis- 
tal cleft on the outer margin) (Figs. 1C, 2C— 
ER Ne 1988) sss 20By 2B Yeo etvalk 
1999: figs. 4A—C, F—H, 7B—E, G—H). 
Johora gua is most similar to J. punicea 
(Ng, 1985), also from Pulau Tioman. Both 
species share a relatively smooth carapace 
dorsal surface, with weakly cristate antero- 
lateral margins, and relatively blunt post- 
orbital cristae, and very low, poorly devel- 
oped epistome posterior margin median 
tooth (Fig. 1A, C; Ng 1988: fig. 22A, B). 
The G1 structure in both species is also 
very similar in having a long, slender out- 
wardly curved or sickle-shaped terminal 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


segment that lacks a dorsal flap, and a rel- 
atively broad subterminal segment that 
lacks a distinct distal cleft on the outer mar- 
gin (see Fig. 2C—E H; Yeo et al. 1999: fig. 
6B—-D). However, J. gua can be distin- 
guished from J. punicea by the distinctly 
slenderer ambulatory legs (e.g., dactylus of 
second leg 11.4—15.0 vs. 5.9—7.5 times lon- 
ger than broad; second ambulatory leg mer- 
us 4.3—4.4 vs. 3.5—4.0 times longer than 
broad) (Fig. 2A, B; Ng & Chong 1986; Ng 
1988: pl. 1 fig. D, fig. 22A); Gl terminal 
segment being less strongly hooked or out- 
wardly curved (vs. Gl terminal segment 
more strongly hooked or curved outwards 
in appearance); and G1 subterminal seg- 
ment being relatively slenderer (vs. G1 sub- 
terminal segment relatively broader) (Fig. 
2C-—E H; Yeo et al. 1999: fig. 6B-D). Fur- 
thermore, specimens of J. punicea are larg- 
er than those of J. gua, with specimens of 
J. punicea (e.g., male, 11.3 X 8.8 mm, ZRC 
1996.1733) similar in size to the holotype 
of J. gua (11.4 .X §8:5 aumieAREe 
2000.2236), possessing undeveloped juve- 
nile Gls (Fig. 2I, J). 

In addition to the above primary differ- 
ences, J. gua can also be immediately sep- 
arated from juvenile as well as adult spec- 
imens of J. punicea by its light orange live 
coloration (vs. deep purple in large adults; 
pink to light purple in juveniles) and pro- 
portionately smaller corneas, about 0.4 
times length of the eyestalks (vs. normal- 
sized corneas, subequal to half length of 
eyestalks) (Fig. 1A, C, D; Ng & Chong 
1986; Ng 1988: pl. 1 fig. D, fig. 22A, B). 
These secondary characters, together with 
the long and slender ambulatory legs seem 
to reflect the cave-dwelling habit of J. gua. 
It is interesting to note here that J. gua is 
more similar to juvenile specimens of J. 
punicea, which have relatively slenderer al- 
beit undeveloped Gls, lighter pigmentation 
and slenderer ambulatory leg segments 
compared to adult specimens, suggesting a 
possible progenetic origin for J. gua from 
J. punicea. 

Johora gua appears to be a true caver- 


VOLUME 114, NUMBER 3 


Fig. 1. A—-C, E-H, Johora gua, new species, holotype male (11.4 X 8.5 mm) (ZRC 2000.2236); D, Johora 
punicea (Ng, 1985), male (11.3 X 8.8 mm) (ZRC 1996.1733). A, dorsal view of carapace; B, left third maxil- 


liped; C, D, frontal view of carapace, left side; E, dorsal view of carpus of right cheliped; EK anterior thoracic 
sternum (sternites 1-4); G, male abdomen; H, posterior thoracic sternum (sternites 5-8). Scales = 2.0 mm (A, 
C, D, F—H); 1.0 mm (B, EB). 


622 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. A—G, Johora gua, new species, holotype male (11.4 X 8.5 mm) (ZRC 2000.2236); H—J, Johora 
punicea (Ng, 1985), H, holotype male (19.7 X 15.5 mm) (ZRC 1984.6803) (after Ng, 1988), I, J, male (11.3 X 
8.8 mm) (ZRC 1996.1733). A, right second ambulatory leg; B, right fourth ambulatory leg; C, I, dorsal view 
of right G1; D, J, ventral view of right G1; E, dorsal view of right G1 terminal segment; E ventral view of right 
G1 terminal segment; G, right G2; H, ventral view of left G1. Scales = 2.0 mm (A, B); 1.0 mm (G, I, J); 0.5 
mm (C—F). 


VOLUME 114, NUMBER 3 


nicolous species, as it has not been found 
in any surface streams on Pulau Tioman de- 
spite extensive and intensive sampling on 
the island over several years (see Yeo et al. 
1999). In addition, J. gua possesses a com- 
bination of troglomorphic features in its 
proportionately long, slender ambulatory 
legs, slightly reduced corneas and reduced 
pigmentation (relative to J. punicea adults). 
Such features are commonly seen in various 
combinations and degrees in troglobitic 
crabs in response to life in a cave habitat, 
where light is either severely reduced or en- 
tirely absent, and the sense of touch be- 
comes more important than sight (see Guin- 
ot 1988; Ng & Sket 1996; Yeo & Ng 1999). 
Johora gua, however, may be an incipient 
troglobite as its troglomorphic features are 
not as highly specialized as that of other 
known troglobitic crabs, such as the potam- 
id Erebusa calobates Yeo & Ng, 1999, 
from Laos, which has much longer and 
slenderer legs together with reduced eye- 
stalks and corneas; the parathelphusid Sun- 
dathelphusa sottoae Ng & Sket, 1986, from 
Philippines, which has very reduced pig- 
mentation as well as corneas; and the hy- 
menosomatid Cancrocaeca xenomorpha 
Ng, 1991, from Sulawesi, which has very 
long, slender legs, and no pigmentation, 
eyes or orbits (see Ng 1991; Ng & Sket 
1996; Yeo & Ng 1999). 


Acknowledgments 


I am grateful to P. K. L. Ng for his con- 
structive criticism of the manuscript; T. M. 
Leong and H. H. Tan for collecting the 
specimens and providing the interesting 
ecological observations; and C. M. Yang 
and K. L. Yeo (ZRC) for access to material 
under their care. 


Literature Cited 


Bott, R. 1966. Potamiden aus Asien (Potamon Savigny 
und Potamiscus Alcock) (Crustacea, Decapo- 
da).—Senckenbergiana biologica 47:469—509. 


623 


Guinot, D. 1988. Les crabes cavernicoles du monde.— 
Mémoires de Biospéologie 15:1—40. 

Ng, P. K. L. 1985. Freshwater decapod crustaceans 
from Pulau Tioman, West Malaysia.—Zoolo- 
gische Mededelingen 59:149—-162. 

. 1988. The freshwater crabs of Peninsular Ma- 

laysia and Singapore. Department of Zoology, 

National University of Singapore, Shinglee 

Press, Singapore, 156 pp., Figs. 1-63. 

. 1990. A new species of Johora Bott, 1966 

(Crustacea: Decapoda: Brachyura: Potamidae) 

from Pulau Redang, Trengganu, Peninsular Ma- 

laysia—Journal of Natural History 24:305— 

310. 

. 1991. Cancrocaeca xenomorpha, new genus 

and species, a blind troglobitic freshwater hy- 

menosomatid (Crustacea: Decapoda: Brachyu- 
ra) from Sulawesi, Indonesia.—Raffles Bulletin 

of Zoology 39:59-73. 

, & S. S. C. Chong. 1986. The freshwater crabs 

and prawns of Pulau Tioman.—Nature Malay- 

siana, Kuala Lumpur 11:26-31. 

, & B. Sket. 1996. The freshwater crab fauna 

(Crustacea: Decapoda: Brachyura) of the Phil- 

ippines. IV. On a collection of Parathelphusidae 

from Bohol.—Proceedings of the Biological So- 

ciety of Washington 109:695—706. 

, & M. Takeda. 1992. On some freshwater 

crabs (Crustacea: Brachyura: Potamidae, Par- 

athelphusidae and Grapsidae) from Peninsular 

Malaysia.—Bulletin of the National Science 

Museum, Tokyo, Series A (Zoology) 18:103— 

116. 

, & L. W. H. Tan. 1984. A new freshwater crab, 
Stoliczia (Johora) tiomanensis n. sp. (Decapo- 
da: Brachyura: Potamidae) from Pulau Tioman, 
West Malaysia.—Malayan Nature Journal 37: 
167-174. 

Ng, H. H., H. H. Tan, & K. K. P. Lim. 1999. The inland 
fishes of Pulau Tioman, Peninsular Malaysia.— 
Raffles Bulletin of Zoology, supplement 6:169— 
187. 

Roux, J. 1936. Second note upon freshwater decapod 
crustaceans from the Malay Peninsula.—Bulle- 
tin of the Raffles Museum, Singapore 12:29— 
33, pls. 12-13. 

Yeo, D. C. J.. & P. K. L. Ng. 1999. Erebusa calobates, 
new genus, new species, a troglobitic crab 
(Brachyura: Potamidae) from Laos.—Journal of 
Crustacean Biology 19:908—916. 

5 WoOX, Cail, Ce IP IG IL, IN, IDQO, WS tives 

water and terrestrial Decapod Crustacea of Pu- 

lau Tioman, Peninsular Malaysia—Raffles Bul- 

letin of Zoology, supplement 6:197—244. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
114(3):624—639. 2001. 


Symbionts of the hermit crab Pagurus longicarpus Say, 1817 


(Decapoda: Anomura): New observations from New Jersey waters 


and a review of all known relationships 


John J. McDermott 


Department of Biology, Franklin and Marshall College, Lancaster, Pennsylvania 17604-3003 


Abstract.—The gastropod shell-Pagurus longicarpus complex harbors a va- 
riety of associates representing different kinds of symbiosis. There are 31 
known relationships, 24 of which have been found with hermit crabs from New 
Jersey, U.S.A. These involve species from eight phyla, living on, embedded 
in, or in the lumen of the gastropod shells, as well as on or in the crab host. 
Crabs primarily inhabited the shells of Nassarius obsoletus (75.7%; n = 6757). 
The hydroid Hydractinia symbiolongicarpus and the bryozoan Alcyonidium 
albescens dominated the outside surfaces of shells (51.5% and 26.5%, respec- 
tively), and a predatory relationship between the bryozoan and the nudibranch 
Corambe obscura was observed. Dipolydora commensalis and Lepidonotus 
sublevis were the most common polychaetes living in shell burrows (38.9%) 
and in the shell lumen (25.8%), respectively. Acrothoracican barnacles, Try- 
petesa lampas, were embedded in 4.7% of shells. In New Jersey P. longicarpus 
is parasitized by the isopods Stegophryxus hyptius and Paguritherium alatum. 
Among known parasites of the crab not yet recorded from New Jersey are the 
isopod Asymmetrione desultor, plerocercoid larvae of the cestode Calliobothrium 
verticillatum, and cystacanth larvae of the acanthocephalan Polymorphus sp. 


The longwrist hermit crab Pagurus lon- 
gicarpus Say, 1817, resides in Atlantic 
coastal waters of the United States from 
Maine to the Gulf of Mexico (Williams 
1984). In the mid-Atlantic region crabs are 
abundant intertidally in depressions on ex- 
posed muddy sand flats during the warmer 
months, but retreat to the subtidal for the 
winter (Rebach 1974, 1978: McDermott, 
pers. obs.). Many aspects of the crab’s bi- 
ology have been studied, including feeding, 
shell acquisition, environmental tolerances, 
reproduction, and symbiotic relationships 
(see McDermott 1999). The term symbiosis 
is used in the original and general way re- 
ferring to “the living together of unlike or- 
ganisms’’ (Committee on Terminology 
IQ37)). 

Pagurus longicarpus harbors a variety of 
symbionts, which are attached to the out- 
side or inside of occupied gastropod shells, 


burrowed into the shell, or free in the shell’s 
lumen, and the crab itself is host to several 
species of ecto and endosymbionts. Some 
of the symbionts are specific for this hermit 
crab, while others, although showing pre- 
dilections for the crab, are found often on 
a variety of other surfaces. Thus, P. longi- 
carpus with its molluscan shell serves as 
host for a diversity of organisms from pro- 
tozoans to crustaceans. 

The present paper documents the species 
of gastropod shells utilized by P. longicar- 
pus and symbionts associated with the 
shells and the crabs themselves from the 
waters of New Jersey. This is followed by 
a review of the biology of all known sym- 
biotic relationships. As stated so colorfully 
by Allee (1923), ““Pagurus longicarpus ap- 
parently roams at will ... carrying with 
him, willy-nilly his commensals.”’ 


VOLUME 114, NUMBER 3 


Materials and Methods 


An intertidal population of Pagurus lon- 
gicarpus located in Hereford Inlet estuary 
(Grassy Sound Channel 39°01.72'N, 
74°48.10'W) has been under study since the 
mid-1960’s (Biggs & McDermott 1973, Ly- 
twyn & McDermott 1976, McDermott 
1998, 1999). Water temperatures range 
from 1° to >25°C, and mean salinity is 
~30%oc. Crabs were collected randomly by 
hand on exposed tidal flats from 1986 to 
1990. They were isolated in the field in 
compartmentalized, plastic boxes contain- 
ing sea water (each of 18 compartments 
were 51 X 43 X 39 mm), or were imme- 
diately preserved in bulk in 10% sea water- 
Formalin and later transferred to 70% ethyl] 
alcohol. The gastropod shells harboring 
hermit crabs were identified, and their 
lengths were measured with Vernier cali- 
pers to 0.1 mm. External symbionts on 
these shells were identified, and those that 
could be removed (e.g., nudibranchs or ca- 
lyptraeid gastropods) were preserved for 
later study. Shells were then cracked with a 
hammer, crabs were isolated in fresh sea 
water, and symbionts embedded in the shell 
fragments or found in the lumen were iden- 
tified. Isolated crabs were sexed and the 
length of the anterior shield of the carapace 
(SL) was measured with Vernier calipers to 
0.05 mm. Preserved crabs were treated in a 
similar manner; however, the prevalence of 
animals not cemented to the shells (e.g., nu- 
dibranchs from the outside of the gastropod 
Shells, and polyclad turbellarians, poly- 
chaetes and calyptraeid gastropods from the 
lumen) could not be determined from the 
preserved collections. Observations on re- 
production were made on some of these 
symbionts. The prevalence of epicaridean 
isopod parasites was determined from live 
and preserved crabs as described in Mc- 
Dermott (1998), although crabs were not 
routinely dissected to look for other internal 
parasites. 


625 
Results 


Of the 6757 shells inhabited by Pagurus 
longicarpus, 95.0% were Nassarius obso- 
letus (Say) and N. trivittatus (Say), 75.7% 
and 19.3%, respectively (Table 1). Crabs 
occupied 137 shells of Urosalpinx cinerea 
(Say) (2.0%), and the other 13 gastropod 
species made up only 3.0%. 

A partially annotated list of all symbionts 
found within the host shell-Pagurus longi- 
carpus complex in this study, as well as all 
other known symbionts, appears in Table 2. 

There were 20 species of invertebrates 
associated with shells harboring P. longi- 
carpus from the live and preserved crab 
collections from New Jersey waters (Table 
3; n = 5041). The hydroid Aydractinia 
symbiolongicarpus and the ctenostome 
bryozoan Alcyonidium albescens were the 
dominant species encrusting outside surfac- 
es of hermit shells, while the polynoid poly- 
chaete Lepidonotus sublevis was the main 
inhabitant of the shell lumen. The two ca- 
lyptraeid gastropods Crepidula convexa and 
C. plana, found on the outside and inside 
of the shell, respectively, were equally 
prevalent. Spat of the blue mussel Mytilus 
edulis was attached to the outside and in- 
side of the shell and the exoskeleton of the 
crab predominately from January to April. 

A prey-predator relationship exists be- 
tween the bryozoan Alcyonidium albescens 
and the nudibranch Corambe obscura. The 
bryozoan was attached to 701 of the 2838 
(24.7%) shells, while the nudibranch and/or 
its coiled egg strings were found on 78 
(2.8%) shells. Three of these shells were 
without the snail itself, but since its egg 
strings were present, it was assumed that at 
least one nudibranch from each shell had 
been dislodged when the crabs were col- 
lected. Except for three shells which lacked 
A. albescens, C. obscura was otherwise 
found only on shells harboring the bryo- 
zoan (96.2%). Only three of the 2137 shells 
without A. albescens (0.14%) had the nu- 
dibranch, while 75 of the 701 shells with 
the bryozoan (10.7%) harbored the preda- 


626 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Gastropod shells inhabited by Pagurus longicarpus in the Hereford Inlet estuary of New Jersey, 
based on all collections from 1985 to 1990 (n = 6757). Arranged in order of decreasing abundance.* 


Species Number Percent 
Nassarius obsoletus (Say) 5112 75.65 
Nassarius trivittatus (Say) 1306 19.33 
Urosalpinx cinerea (Say) 137 2AOB) 
Neverita (= Polinices) duplicata (Say) 59 0.87 
Eupleura caudata (Say) 45 0.67 
Busycon carica (Gmelin) 23 0.34 
Littorina littorea (Linnaeus) 2 0.31 
Euspira (= Lunatia) heros (Say) 19 0.28 
Littoraria (= Littorina) irrorata (Say) 12 0.18 
Busycotypus (= Busycon) canaliculatus (Linnaeus) 9 0.13 
Costoanachis (= Anachis) avara (Say) 7 0.10 
Astyris (=Mitrella) lunata (Say) 3 0.04 
Epitonium rupicola (Kurtz) 2) 0.03 
Littorina obtusata (Linnaeus) 1 0.02 
Nassarius vibex (Say) 1 0.02 


* Classification based on American Fisheries Society (1998). 


tor. Thus, C. obscura was 76.4 times more 
abundant on shells with this bryozoan. 

Ninety-nine specimens of C. obscura 
were recorded from 78 shells (1—4/shell), 
and egg strings were found on 17 shells (1— 
7/shell). The maximum number of seven 
strings occurred with a single nudibranch. 
Observations suggested that there are two 
periods of reproduction, April-May and 
September—November. Veligers hatched 
only from eggs collected in October and 
November. Mean length of living nudi- 
branchs was 2.41 + 1.49 mm, range 0.69— 
4.40 mm (n = 14), and snails approximate- 
ly 23.0 mm were mature. One circular egg 
string measured 2.3 mm in diameter. 

The spionid polychaete Dipolydora com- 
mensalis was the dominant species that 
bores into the shell, having a prevalence ap- 
proximately eight times that of the acro- 
thoracican barnacle Trypetesa lampas. The 
prevalence of these symbionts was com- 
pared between the two most common host 
shells, Nassarius obsoletus and N. trivitta- 
tus. Dipolydora commensalis occurred in 
761 of 2110 (36.1%) of N. obsoletus shells 
and 243 of 570 (42.6%) of N. trivittatus 
shells (significant difference, contingency 
x? = 8.28, p = <0.005). The worm also 
occurred in shells of six of the other gas- 


tropod species listed in Table 1 (31 of 157, 
19.8%). Worms were reproducing most of 
the time that crabs were on the intertidal 
flats, but no data are available while crabs 
were subtidal during the winter (December— 
February). Female worms with mature ova, 
along with their egg cases in the burrows 
were seen as early as March, and 3-setiger 
larvae were hatching in April when the wa- 
ter temperatures during these months 
ranged from approximately 7° to 12°C. Re- 
cently settled juveniles appeared to be most 
common in the fall. Numbers of worms per 
shell were usually not counted, but a max- 
imum of eight worms were found in a sin- 
gle burrow, only one of which was a mature 
female. 

Trypetesa lampas was found in 66 of 
2110 (3.1%) N. obsoletus shells and 9 of 
571 (1.6%) N. trivittatus shells (significant 
difference, contingency x? = 4.01, p = 
<0.05). A maximum of four barnacles were 
found in N. obsoletus shells, while no more 
than one occurred in N. trivittatus. Unlike 
D. commensalis, T. lampas was not found 
in 157 of the other shell species. Barnacles 
with eggs and developing nauplii were ob- 
served in August and September of 1987, 
and liberated nauplii were seen in August 
1988. 


VOLUME 114, NUMBER 3 


Epicaridean symbionts on and in the her- 
mit crabs from the same population were 
studied recently (McDermott 1998) and are 
listed in Table 2. Their prevalence and bi- 
ology, along with other crab parasites (Ta- 
ble 3) not yet found in New Jersey waters 
(Table 2) are discussed below. 


Discussion 


Crab shell utilization (Yable 1).—The 
species of gastropod shells used by P. lon- 
gicarpus varies over the latter’s geographi- 
cal range. In waters north of New Jersey 
(Connecticut and Rhode Island) the shells 
of Littorina littorea are used in numbers 
equal to those of Nassarius obsoletus 
(~25%) (Blackstone & Joslyn 1984), or 
greatly outnumber the latter by ~80% to 
<5%) (Scully 1979). The habitat of N. ob- 
soletus in New England has gradually con- 
tracted due to competitive exclusion since 
the introduction of L. littorea from Europe 
in the mid-1880’s (Brenchley & Carlton 
1983). New Jersey is near the southern limit 
of L. littorea, and its populations are rela- 
tively small. Shells of the intertidal gastro- 
pods Littorina saxatilis (Olivi) and Nucella 
lapillus (Linnaeus), species not found in 
New Jersey, are inhabited infrequently by 
P. longicarpus in Rhode Island (Scully 
1979). At Beaufort, North Carolina, where 
L. littorea is rare, N. obsoletus and the more 
southern N. vibex are utilized in equal num- 
bers (~30%) (Kellogg 1977). In the same 
area Costoanachis avara, Terebra dislocata 
(Say) and Urosalpinx cinerea are each rep- 
resented at ~8%. In the North Inlet estuary 
of South Carolina, Young (1979) found that 
56.5% of 1208 P. longicarpus were in Nas- 
sarius spp. shells (he did not distinguish the 
species, but the majority were probably WN. 
vibex). Terebra dislocata and U. cinerea 
were each used by ~15% of the crabs. In 
the northeastern Gulf of Mexico (Florida) 
Littoraria irrorata and N. vibex are inhab- 
ited by 80% of the P. longicarpus popula- 
tion (Kuhlmann 1992). In the Galveston 
area of the Texas coast, Fotheringham 


627 


(1976) recorded 47% of the crabs in shells 
of Neverita duplicata and ~15% each in L. 
irrorata and Stramonita (=Thais) haemas- 
toma floridana (Conrad). Littoraria irrora- 
ta, the southern periwinkle, is rarely found 
in New Jersey and northward (Blackstone 
1986, McDermott, pers. obs.). In all of the 
cases mentioned above, it is primarily the 
Shell of the dominant gastropods in the re- 
gion that are used by P. longicarpus. As 
Fotheringham (1976) and others have not- 
ed, shell-use patterns are influenced by a 
variety of factors including: abundance of 
shells, shell size, internal volume and shell 
morphology, crab ontogeny, sex, intra and 
interspecific competition, and attached sym- 
bionts. Besides the 15 gastropod shells in 
Table 1 and Terebra dislocata and Stra- 
monita haemastoma floridana, Sumner et 
al. (1913) noted that Euspira triseriata 
(Say), E. immaculata (Totten) and Colus 
pygmaeus (Gould) were also used by P. 
longicarpus. Thus, P. longicarpus has been 
shown to utilize a total of 20 different gas- 
tropod species along its range on the east 
coast. 

Symbionts associated with the shell (Ya- 
bles 2, 3).—The colonial hydroids Hydrac- 
tinia symbiolongicarpus Buss & Yund and 
Podocoryne carnea Sars are both faculta- 
tive symbionts. The former has recently 
been recognized as one of three sibling spe- 
cles associated with P. longicarpus. It is 
distinguished from H. symbiopollicaris 
Buss & Yund, H. polyclina (Agassiz) and 
the European H. echinata (Fleming) by 
breeding incompatabilities, electrophoretic 
analyses, morphometrics, and some speci- 
ficity for crab hosts (Buss & Yund 1989). 
H. symbiolongicarpus and H. symbiopolli- 
caris are found mainly on the shells occu- 
pied by P. longicarpus and P. pollicaris 
Say, respectively, but the reverse sometimes 
occurs. Originally H. polyclina was found 
only on shells inhabited by Pagurus aca- 
dianus Benedict, and not on P. longicarpus 
or P. pollicaris (Buss & Yund 1989), but a 
more recent study has shown that H. poly- 
clina may also be found on shells harboring 


628 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Master list of symbionts of Pagurus longicarpus arranged according to their location in the host 
shell-hermit crab complex and in phylogenetic order within the location. Most were found in crabs from New 


Jersey waters. 


Symbionts 


Remarks* 


On Outside of Shell 
Cnidaria 


Hydractinia symbiolongicarpus Buss & Yund 


Podocoryne carnea Sars 


Ectoprocta 
Alcyonidium albescens Winston & Key 
Conopium tenuissimum (Canu) 
Membranipora tenuis Desor 
Schizoporella unicornis (Johnston) 
Annelida 


Hydroides dianthus (Verrill) 
Sabellaria vulgaris (Verrill) 


Mollusca** 


Crepidula convexa Say 

Crepidula fornicata (Linnaeus) 

Cuthona (nana ?) (Alder & Hancock) 
Corambe (=Doridella) obscura (Verrill) 


Mytilus edulis Linnaeus 


Arthropoda 


Anoplodactylus (lentus) Wilson 
Balanus eburneus Gould 
Balanus improvisus Darwin 


Bored into Shell 
Annelida 


Dipolydora (=Polydora) commensalis (An- 
drews) 

Dipolydora socialis (Schmarda)*** 

Polydora neocaeca Williams and Radashey- 
Shayari 


Polydora websteri Hartman*** 


Arthropoda 
Trypetesa lampas (Hancock) 
Lumen of Shell 
Platyhelminthes 
Stylochus zebra (Verrill) 
Annelida 


Lepidonotus sublevis Verrill 


Mollusca 
Crepidula plana Say 
External on Crab 
Protozoa 


Lagenophrys eupagurus Kellicott*** 
Unidentified folluculinid ciliate 


One of three sibling species associated with crabs in 
genus Pagurus (see Discussion) 

Shows a predilection for the shells of Nassarius trivit- 
tatus 


Facultative symbiont 
Facultative symbiont 
Facultative symbiont 
Facultative symbiont 


Facultative symbiont 
Facultative symbiont 


Shows some predilection for hermit shells 

Seldom with this crab, often on P. pollicaris 

Predator of Hydractinia 

Predator of Alcyonidium and other encrusting bryozo- 
ans 

Seasonal symbiont as spat in breeding seasons (Janu- 
ary to April and June) 


Found among the polyps of Hydractinia 
Facultative symbiont 
Facultative symbiont 


Bores into columella, and may make contact with shell 
lumen; obligate symbiont 

Superficial burrow on outside of shell 

Lives in an unbranched U-shaped burrow in hermit 
shells as well as living gastropods and bivalve shell 
fragments 

Superficial burrow on outside of shell 


Bores from lumen side of body whorl 


Embryo predator of P. pollicaris 


Obligate symbiont 


Shows predilection for hermit shells 


Tests containing peritrichous ciliate attached to gills 
Tests containing heterotrichous individuals cemented to 
body 


VOLUME 114, NUMBER 3 


Table 2.—Continued. 


Symbionts 


629 


Remarks* 


Mollusca 


Mytilus edulis Linnaeus 


Arthropoda 


Stegophryxus hyptius Thompson 


Internal in Crab 
Platyhelminthes 


Calliobothrium verticillatum (Rudolphi)*** 


Acanthocephala 


Polymorphus sp.*** 


Arthropoda 


Paguritherium alatum Reinhard 


* Additional information in Discussion. 


Seasonal symbiont 


Bopyrid isopod in branchial chambers 
Bopyrid isopod on abdomem 


Plerocercoids of tetraphyllidean cestode in Pagurus 
pollicaris, but possibly also in P. acadianus and P. 
longicarpus 


Cystacanths in abdominal hemocoel 


Entoniscid isopod in hemocoel 


** Mollusk classification based on American Fisheries Society (1998). 


«= Not identified from New Jersey crabs. 


P. longicarpus (Folino & Yund 1998). Hy- 
dractinia from New Jersey is assumed to 
be A. symbiolongicarpus. The high preva- 
lence of Hydractinia on hermit shells 
(>50%) in New Jersey is consistent with 
Lytwyn’s (1979) observations in the same 
State in the 1970’s. In Texas waters its prev- 
alence was approximately 30% (Fother- 
ingham 1976). 

The prevalence of Podocoryne carnea on 
hermit shells from New Jersey was much 
lower (0.5% of 2838 crabs) than for H. 
symbiolongicarpus (TYable 3). McFadden 
(1986) found that P. carnea was rare 
among the epifauna of P. longicarpus shells 
collected in Long Island Sound. She also 
noted that recruits of this hydroid were 
found on only 0.2% of 1663 shells but did 
not record the species of gastropod shells 
that harbored P. carnea. All 15 colonies of 
this species from New Jersey crabs were on 
the shells of Nassarius trivittatus, which 
agrees with Crowell (1945) who showed 
that P. carnea has a predilection for the 
crab-occupied shells of N. trivittatus, al- 
most never being found on the shells of Lit- 
torina littorea, at least in the Woods Hole 


region of Massachusetts. He found both P. 
carnea and Hydractinia sp. on crab-occu- 
pied shells of Busycon sp., Urosalpinx ci- 
nerea, Eupleura caudata, Nassarius obso- 
letus, and others. Unlike Crowell, Edwards 
(1972) identified colonies of P. carnea on 
Littorina shells occupied by hermit crabs 
from the British Isles. Podocoryne carnea 
has free medusae in its life cycle (unlike 
Hydractinia), which are liberated from gon- 
ozooids whose white color makes the col- 
ony easily recognizable. Gonozooids may 
be seasonal. Medusae develop and are lib- 
erated from late May to early August in col- 
onies on New Jersey crabs. The presence of 
P. longicarpus inside of a Nassarius trivit- 
tatus shell covered with P. carnea in some 
manner allows for the differentiation of spi- 
ral zooids around the periphery of the ap- 
erture (Braverman 1960). Although the 
mechanism involved in this differentiation 
and the functional significance of this phe- 
nomenon have not been explained, it is pos- 
sible that these elongated zooids may be 
utilized for capturing food items stirred up 
from the sediments by the hermit crab or 
crab zoeae liberated from the shell aperture. 


630 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 3.—Prevalence of symbionts found on, in, or in the lumen of gastropod shells inhabited by Pagurus 
longicarpus from the Hereford Inlet estuary of New Jersey 1986—1990, based on the examination of shells with 
live (n = 2838) and preserved crabs (n = 2203). Species within each phylum are arranged alphabetically. 


Live crabs Preserved crabs 
Species Number Percent Number Percent 
Cnidaria 
Hydractinia spp. 1475 52.0 1120 50.8 
Podocoryne carnea 15 0.5 ND* 
Ectoprocta 
Alcyonidium albescens 701 24.7 634 28.8 
Other encrusting species** 35 1.2 76 3.4 
Platyhelminthes 
Stylochus zebra 2 0.1 ND 
Annellida 
Dipolydora commensalis 1038 36.6 Sy 41.9 
Hydroides dianthus 10 0.4 6 0.3 
Lepidonotus sublevis 612 21.6 688 By 
Sabellaria vulgaris 90 Be 47 esl 
Mollusca 
Corambe obscura 78 2.8 ND 
Crepidula convexa 318 11.2 ND 
Crepidula fornicata 2 0.1 ND 
Crepidula plana 320 11.3 ND 
Mytilus edulis spat ZIN3) 7.5 ND 
Pycnogonida 
Anoplodactylus (lentus) 4 0.1+ 
Arthropoda 
Barnacle spat*** 34 Ee) ND 
Trypetesa lampas 75 2.6 163 7A 


* ND = no data. 


** Conopium tenuissimum (Canu), Membranipora tenuis Desor, Schizoporella unicornis (Johnston). 


*** Balanus eburneus and B. improvisus. 


Additional invertebrates that are only loosely associated with shells, primarily in silt accumulated in damaged 
apices: sea anemone; heteronemerteans, hoplonemerteans; polychaetes, Polycirrus eximus (Leidy), Autolytus sp., 
phyllodocids, spionids; amphipods, Caprella sp., Corophium sp., isopod, Erichsonella filiformis (Say); mollusc, 
Anadara ovalis (Bruguiére). Living boring sponge Cliona sp. was found on one Nassarius obsoletus shell. 


Pagurus longicarpus and P. pollicaris seem 
to prefer gastropod shells with Hydractinia 
colonies, while some other members of the 
genus and other genera may reject shells 
with this hydroid (Conover 1976, Mills 
1976, Mercando & Lytle 1980). Other ev- 
idence suggests that P. longicarpus does 
not discriminate between shells with or 
without HA. symbiolongicarpus Weissberger 
15). 

The only turbellarian associated with P. 
longicarpus 1s the polyclad Stylochus zebra, 


but it is much more commonly found in the 
lumen of shells inhabited by P. pollicaris 
(Lytwyn 1976, 1979; Lytwyn & Mc- 
Dermott 1976). Its rarity with P. longicar- 
pus in the present study confirms that found 
previously in New Jersey by Lytwyn & 
McDermott (1976) and Lytwyn (1979). 
This polyclad has been recorded from three 
other species of hermit crabs from North 
Carolina [Pagurus impressus (Benedict), 
Petrochirus diogenes (Linnaeus) and Dar- 
danus venosus (H. Milne Edwards)], all 


VOLUME 114, NUMBER 3 


with a lower prevalence than with P. pol- 
licaris (Lytwyn 1979). It is an embryo 
predator of P. pollicaris (Lytwyn 1979), but 
also feeds on shell symbionts such as the 
white slipper limpet Crepidula plana (Ly- 
twyn & McDermott 1976). 

For many years the encrusting ctenosto- 
me bryozoan Alcyonidium albescens was 
known as A. polyoum (Hassall). However, 
Winston & Key (1999) have shown recently 
that the latter is a European species. Al- 
cyonidium albescens is a common faculta- 
tive inhabitant of the outer surfaces of P. 
longicarpus-inhabited Nassarius obsoletus 
shells, but is rarely associated with the 
shells of the living snail (Karlson & Car- 
iolou 1982). It grows, however, on the 
shells of some other living snails and on 
shells occupied by P. pollicaris, as well as 
on brachyuran crabs and inanimate shell 
surfaces (Karlson & Cariolou 1982, Karl- 
son & Shenk 1983, Key et al. 1999, 
McDermott pers. obs.). Buss & Yund 
(1988) found that it dominates Hydractinia 
Spp. in competition for space on shells oc- 
cupied by P. longicarpus. 

The lumen of shells with P. longicarpus 
(and P. pollicaris) often contains the obli- 
gate polynoid polychaete symbiont, Lepi- 
donotus sublevis Verrill (Pettibone 1963, 
Lytwyn 1979, Mercando 1983, McDermott 
pers. obs.). It was found in the lumen of 
approximately 25% of hermit crab shells in 
New Jersey (Table 3). This was about half 
of that recorded by Lytwyn (1979) in the 
1970’s from a much smaller sample. Lepi- 
donotus sublevis was found to be 25 times 
more frequent in shells harboring male 
crabs, which may be related to crab size 
(males tend to be larger than females) rather 
than to sex per se (Mercando 1983). These 
scaled worms may occupy a considerable 
percentage of the shell’s internal volume, 
and Mercando & Donaghy (1984) showed 
that P. longicarpus had more difficulty en- 
tering shells occupied by L. sublevis, 1.e., 
they selected empty shells significantly 
more frequently than those with worms. Al- 
though Frotheringham (1976) demonstrated 


631 


that L. sublevis consumed detached embry- 
os of P. longicarpus in the laboratory, there 
is still no evidence that the worm is an em- 
bryo predator under natural conditions. 

Five species of gastropods are associated 
with P. longicarpus (Table 2). Of the three 
slipper limpets (Calyptraeidae), Crepidula 
fornicata (Linnaeus), C. plana Say and C. 
convexa Say, only the last two seem to 
show some predilection for inhabiting P. 
longicarpus shells (Table 3) (Karlson & 
Cariolou 1982, McDermott pers. obs.). C. 
plana lives on the inside of the shells, while 
C. convexa and C. fornicata are on the out- 
side. The geographical distribution of C. 
plana is from New England to Georgia, and 
it is distinct from C. depressa Say, an in- 
habitant of hermit crab shells from Florida 
to Texas (Collin, 2000). Crepidula plana 
has a distinct relationship with crab-inhab- 
ited shells and is rarely detected on other 
substrates. This applies to shells inhabited 
by P. longicarpus (Shenk & Karlson 1986, 
Table 3), as well as P. pollicaris (Shenk & 
Karlson 1986, McGee & Targett 1989). Al- 
though the veligers of C. plana may settle 
and metamorphose on a variety of sub- 
strates, including the outside of hermit crab 
shells, they are usually found only on the 
inside of these shells where they are more 
protected from predation (Shenk & Karlson 
1986, Collin 2000, present study). In this 
location, however, the limpet may be vul- 
nerable to predation if the polyclad Stylo- 
chus zebra is present (Lytwyn & Mc- 
Dermott 1976, Lytwyn 1979). Effluents 
from C. plana and P. pollicaris in Busycon 
carica shells attract competent larvae to set- 
tle and metamorphose (McGee & Targett 
1989). 

The shell morphology of C. convexa liv- 
ing on hermit shells is markedly different 
in its dimensions than those on other sub- 
strates, i.e., the shells are shorter and nar- 
rower but greater in height (Franz & Hen- 
dler 1970). Unlike C. fornicata and C. pla- 
na which release veliger larvae, C. convexa 
has direct development and thus has a 
greater problem in distributing juveniles to 


632 


new locations. Attachment of C. convexa to 
the mobile hermit crab along with the high 
mobility of juvenile snails, however, aids in 
the colonization of new substrates (Hendler 
& Franz 1971). Thus, this association is a 
distinct benefit to C. convexa populations. 
Two nudibranchs are associated with the 
gastropod shell of P. longicarpus. Strong 
evidence is presented here that the dorida- 
cean Corambe obscura has a very close as- 
sociation with one of its prey species, the 
encrusting bryozoan Alcyonidium albes- 
cens. Because of the possibility that some 
nudibranchs were dislodged from _ shells 
while being isolated in the field, it is likely 
that their 10.7% prevalence on the bryo- 
zoan is a minimal value. In Delaware Bay, 
New Jersey, Franz (1967) found that this 
nudibranch was always associated with, and 
fed on, the following encrusting bryozoans: 
Alcyonidium verrilli Osburn, Conopeum 
tenuissimum and Membranipora tenuis 
(Dudley (1973) clarified the identity of the 
last two species). Corambe obscura was 
also found with a A. albescens colony on a 
Neverita duplicata shell harboring Pagurus 
pollicaris from the same New Jersey loca- 
tion (McDermott, pers. obs.). Karlson & 
Shenk (1983) did not recover C. obscura 
from A. albescens colonies living on 39 
shells of Busycon carica with Pagurus pol- 
licaris and 18 live snails collected at the 
mouth of Delaware Bay. Alcyonidium ver- 
rilli, an upright species capable of forming 
large masses, has not been identified on 
Shells with P. longicarpus. Cory (1967) 
found that C. obscura fed and deposited its 
egg strings on the colonies of C. tenuissi- 
mum and M. tenuis in the upper part of the 
Chesapeake Bay, Maryland. Wass (1972) 
reported that this snail is “often abundant 
on Alcyonidium” (presumably referring to 
A. verrilli) in the lower Chesapeake Bay, 
Virginia. Perron & Turner (1977) found the 
coiled egg strings of the snail on its chei- 
lostome prey C. tenuissimum, and also not- 
ed that its larvae metamorphose on the col- 
ony. Perhaps an obligate metamorphosis 
may be involved in the C. obscura—A. al- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


bescens relationship. The geographic distri- 
bution of C. obscura ranges in the Atlantic 
Ocean from the eastern United States to 
Brazil, and is found along the coast of 
northwestern Europe. It was introduced into 
the Black Sea in the late 1980’s (Rogin- 
skaya & Grintsov 1990), and has been 
found subsequently in many locations with- 
in the Sea where it has apparently found 
encrusting bryozoans to sustain its repro- 
duction (Roginskaya & Grintsov 1995, 
oT): 

Cuthona nana (Alder & Hancock) is a 
predator of Hydractinia polyclina, which is 
found primarily on shells inhabited by Pa- 
gurus acadianus in New England (Harris et 
al. 1975, Rivest 1978, Lambert 1991, Fol- 
ino 1987, 1993, 1997). An unidentified spe- 
cies belonging to the same genus was found 
on Hydractinia symbiolongicarpus colonies 
on three shells of Nassarius obsoletus oc- 
cupied by P. longicarpus in April and May 
1986, from the same New Jersey location 
but not part of the collections in Table 3. 
Egg strings of the snail were on two of the 
shells, and on one shell with two nudi- 
branchs most of the hydroid colony was de- 
stroyed. Lengths of the living nudibranchs 
were 3.5 mm for a juvenile and 10, 12 and 
13 mm for adults. 

Four shells among 2838 (0.1%) with liv- 
ing crabs had pycnogonids associated with 
colonies of Hydractinia (July and August 
1987, May 1988). Three of the shells were 
Nassarius trivittatus and the other was N. 
obsoletus. The maximum number per shell 
was five (total = 10). These symbionts were 
juveniles and belonged to the genus Ano- 
plodactylus (probably A. lentus Wilson). 
Anoplodactylus lentus is a common species 
along the eastern coast of the United States, 
living among and feeding on hydroids 
(Cole 1906, Sumner et al. 1913, Hedgpeth 
IQ50, 

Endolithic species (Tables 2, 3).—The 
shell-boring, spionid polychaete Dipolydo- 
ra commensalis is an obligate commensal 
of gastropod shells occupied by hermit 
crabs including: Pagurus annulipes (Stimp- 


VOLUME 114, NUMBER 3 


son), P. longicarpus, P. pollicaris and Cli- 
banarius vittatus Bosc (Andrews 1891, 
Hatfield 1965, Radashevsky 1989, Dauer 
1991). It occurred in approximately 40% of 
the shells inhabited by P. longicarpus in 
this study, which compares well with the 
36.8% prevalence (186 of 506) in five spe- 
cies of shells occupied by four species of 
hermit crabs, 91.8% of which were P. lon- 
gicarpus, in Virginia (Dauer 1991). The 
other species of gastropod shells bored by 
D. commensalis from New Jersey were 
Urosalpinx cinerea, Eupleura caudata, 
Neverita duplicata, Littoraria irrorata, Lit- 
torina littorea, and Busycotypus canalicu- 
latus. Hatfield (1965) recorded the worm 
from the shells of Euspira heros and Buc- 
cinum undatum Linnaeus. Although the 
prevalence of D. commensalis was signifi- 
cantly greater in the shells of Nassarius tri- 
vittatus than N. obsoletus, | have no ratio- 
nale to explain this difference, except for 
possible dissimilarities in mineralization 
that may make the one shell more favorable 
for boring. The polychaete bores into the 
columella of shells producing a tube which 
in some cases eventually enters the lumen 
of the shell near the apex (Andrews 1891, 
Hatfield 1965, Radashevsky 1989, Dauer 
1991, Williams 1995). The anterior end of 
the worm is usually located at the opening 
of the tube in the columella which is locat- 
ed on the medial side of the shell aperture. 
Here the worm extends its palps and draws 
food particles into the mouth (Dauer 1991, 
Williams & McDermott 1997). J. D. Wil- 
liams (per. comm.) found that worms whose 
burrows reach the shell lumen, prey on the 
developing embryos attached to the pleo- 
pods of P. longicarpus in Rhode Island. 
Williams (1999, 2000, 2001) also reported 
that Polydora spp. and Trypetesa sp. inhab- 
iting gastropod shells with hermit crabs 
from the Indo-West Pacific are also embryo 
predators, so that this may be a more com- 
mon phenomenon than previously recog- 
nized. Pagurus longicarpus living in shells 
bored by D. commensalis, rather than nor- 
mal shells, may be more vulnerable to pre- 


633 


dation because these shells are less resistant 
to crushing forces (Buckley & Ebersole 
1994). 

Other spionid polychaetes are boring fac- 
ultative symbionts of shells inhabited by P. 
longicarpus. Polydora websteri Hartman 
was reported in shells from Maine (Blake 
& Evans 1973), and P. neocaeca, recently 
described by Williams & Radashevsky 
(1999), was recovered from shells in Rhode 
Island. Also in shells from this state was 
Dipolydora socialis (Schmarda) (Williams 
per. comm.). The superficial nature of the 
burrows of these species, however, suggests 
that they are not embryo predators. 

The acrothoracican barnacle Trypetesa 
lampas burrows into gastropod shells oc- 
cupied by hermit crabs (Tomlinson 1969a, 
1969b), and was detected in approximately 
5% of the 5041 shells examined in New 
Jersey. The significant difference in the 
prevalence of this barnacle between Nas- 
sarius obsoletus and N. trivittatus may be 
related to the larger size attained by the for- 
mer, some difference in the calcereous com- 
position of the shells, or the much greater 
abundance of N. obsoletus in the Hereford 
Inlet estuary. Barnacle burrows are detected 
only on the inner surfaces of the body 
whorl of cracked shells where slit-like 
openings allows the cirri to protrude into 
the lumen. Beyond this opening is the en- 
larged chamber harboring the barnacle. 
Other acrothoracican barnacles burrow into 
a variety of calcareous substrata (e.g., gas- 
tropod and bivalve shells, corals), but T. 
lampas is an obligate associate of hermit 
crabs. This association obviates harmful 
silting experienced by species living in un- 
inhabited gastropod or bivalve shells (Tom- 
linson 1969a, 1969b). In New England wa- 
ters, this barnacle is found in the shells of 
Euspira heros and Neverita duplicatus oc- 
cupied by hermit crabs (Sumner et al. 1913, 
Zullo 1963). Neverita duplicatus shells in- 
habited by Pagurus pollicaris from New 
Jersey are also infested with T. lampas 
(McDermott pers. obs.). White (1969) 
found T. lampas in the shells of several spe- 


634 


cies of gastropods inhabited by Pagurus 
bernhardus (Linnaeus) in Wales. Some spe- 
cies of shells were heavily infested, e.g., 
67.2% of 2249 Buccinum undatum Linnae- 
us shells were infested, and there were pos- 
itive correlations between shell length and 
the prevalence and numbers per shell 
(White 1969). 
Tomlinson (1969a) said that 7. lampas 
. in general does little if any harm to 
the host. All species of the order collect 
food without taking from or giving any- 
thing of value to the host,” although the 
gastropod shell may be weakened by the 
presence of numerous barnacles. Recent 
studies by Williams (1999) may eventually 
modify Tomlinson’s observations. He 
showed that a species of 7rypetesa found 
in hermit crab shells from the Philippine Is- 
lands ingest crab embryos in nature. 
Found on or in the crab (Tables 2, 3).— 
Two species of ciliated protozoans are ec- 
tosymbionts of P. longicarpus. One is the 
loricate peritrich Lagenophrys eupagurus 
Kellicott, 1893, (Lagenophryidae) that at- 
taches to the crab’s gills (Kellicott 1893, 
Clamp 1989, Fernandez-Leborans & Tato- 
Porto 2000). It is not specific for P. longi- 
carpus, having been reported from thirteen 
other species of crustaceans, none of which 
was a hermit crab (Clamp 1989). Members 
of this genus are well-known as ectosym- 
bionts of crustaceans. The gill filaments of 
hermits from New Jersey were not exam- 
ined for this species. The unidentified cili- 
ate (Table 2) is a heterotrichous folliculinid 
that was periodically found cemented to 
different parts of the crab’s body and also 
on crab shells. The species appears to be a 
facultative commensal and is similar to Pla- 
tyfolliculina paguri Andrews & Reinhard, 
1943, which attaches to the body of Pagu- 
rus pubescens Kroyer, from the coast of 
Maine (Andrews & Reinhard 1943). 
Plerocercoids of the tetraphyllidean ces- 
tode, Calliobothrium verticillatum (Rudol- 
phi) are found in the tubules of the anterior 
midgut ceca of Pagurus pollicaris Say col- 
lected in the Woods Hole region of Mas- 


ce 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


sachusetts (Caira & Ruhnke 1991). Cherry 
et al. (1991) suggested that >95% of hermit 
crabs from this area were infected (it is not 
clear whether this value referred just to P. 
pollicaris or all three hermits from the re- 
gion). Smolowitz et al. (1993) demonstrated 
in histological sections that these plerocer- 
coids cause inflammation to the mid gut 
ceca of Pagurus spp. (Pagurus acadianus, 
P. longicarpus and P. pollicaris). Unfortu- 
nately, although 39 crabs were sectioned for 
study, the authors did not record the identity 
of the crabs (R. M. Smolowitz, pers. 
comm.). The adult parasite is found in the 
spiral valve of the smooth dogfish Mustelus 
canis (Mitchill) (Caira & Ruhnke 1991). 
The occurrence of Mytilus edulis spat at- 
tached to the body of P. longicarpus on a 
seasonal basis is not unprecedented, be- 
cause such was reported more than a half 
century ago from Woods Hole, Massachu- 
setts by Andrews & Reinhard (1943). 
Pagurus longicarpus serves as an inter- 
mediate host for the acanthocephalan Po- 
lymorphus sp. in the Woods Hole area 
(Reinhard 1944). Cystacanths were found 
in the abdominal cavity on the outer walls 
of the gut or among the tubules of the di- 
gestive gland. Usually there was only one 
cystacanth per host, and approximately one 
percent of the crabs were infected. These 
cysts are glistening white, have a mean 
length and width of 2.6 and 1.3 mm, and 
are easily seen through the thin abdominal 
wall (Reinhard 1944). During the course of 
the present study only a small but unre- 
corded percentage of the live crabs were 
dissected, and no cystacanths were found. 
Pagurus longicarpus harbors three epi- 
caridean isopod parasites, two belonging to 
the Bopyridae and the other to the Enton- 
iscidae (McDermott 1998). One of the bo- 
pyrids, Stegophryxus hyptius Thompson 
(subfamily Althelginae), lives on the ab- 
domen of the crab. At the beginning of the 
past century it had a prevalence of 1.5— 
4.0% in the Woods Hole region of Massa- 
chusetts (Thompson 1902), and at mid-cen- 
tury Reinhard (1943, 1949) found the iso- 


VOLUME 114, NUMBER 3 


pods in 1.25—1.5% of the crabs examined. 
During the last part of the century crabs 
from the waters of New Jersey had a prev- 
alence of 0.15% (14 of 9111) (McDermott 
1998). This parasite is found at least as far 
south as Georgia and has little host speci- 
ficity, parasitizing at least seven other spe- 
cies of related hermits (Markham 1974, 
1988). Stegophryxus hyptius does not pro- 
duce any noticeable changes in primary or 
secondary (pleopods) sex characters in P. 
longicarpus (Yhompson 1902, Reinhard et 
al. (1947). 

The other bopyrid, Asymmetrione desul- 
tor Markham, 1975 (subfamily Pseudioni- 
nae), is found along the southeastern coast 
of the United States, inhabiting the bran- 
chial chambers of P. longicarpus (Mark- 
ham 1975). This parasite has been found on 
five other species of hermits (Markham 
1988), but it was not seen in the hermits 
from New Jersey. 

The hemocoel-dwelling entoniscid Pa- 
guritherium alatum Reinhard, 1945, was re- 
covered from 38 of 4600 (0.8%) hermits 
collected in 1943 in the Woods Hole area 
(Reinhard 1943, 1945). In 1946, Reinhard 
& Buckeridge (1950) collected ~100 in- 
fected P. longicarpus from the same locale, 
for the purpose of documenting the effect 
of the parasites on the secondary sex char- 
acters of the hosts, but prevalence was not 
recorded. Adkinson and Heard (1978) 
found the entoniscids in 3% of ~300 crabs 
that they examined in North Carolina. An 
alternate host for P. alatum in North Car- 
olina is Pagurus annulipes; two of ~250 
crabs were parasitized (Adkinson & Heard 
1978). In New Jersey the prevalence of P. 
alatum was only 0.11% (4 of 3703). Pa- 
guritherium alatum castrates its female host 
and causes a pronounced reduction in the 
length of endopodal rami and a reduction 
of endopodal hairs (Reinhard & Buckeridge 
1950); the parasitized male is not modified 
externally. Simultaneous parasitism in P. 
longicarpus with P. alatum and Stegophry- 
xus hyptius occurs, but is rare (Reinhard & 
Buckeridge 1950). 


635 


Rhizocephalans have never been reported 
from Pagurus longicarpus, but its congener 
P. pubescens Kroyer, 1838, from the waters 
of Maine harbors Peltogaster paguri Rath- 
ke (prevalence up to 25%), which also par- 
asitizes the European P. bernhardus (Rein- 
hard 1939, 1942, Walker & Pearse 1939). 
Pagurus acadianus Benedict, from the 
same locality as P. pubescens is not para- 
sitized by this barnacle. 


Conclusion 


Within this gastropod shell-Pagurus lon- 
gicarpus complex are a number of biolog- 
ical relationships that deserve experimental 
study to further define their nature. This ap- 
plies also to other species of hermit crabs 
whose associates are well known, e.g., the 
European P. bernhardus (Jensen & Bender 
1973, Lancaster 1988). Such relationships 
involve a variety of predators and prey both 
on the outside and in the lumen of the 
Shells, shell symbionts involved as crab em- 
bryo predators (a type of periodic parasit- 
ism involving only crabs of one sex), and 
host-parasite relationships involving the 
crabs as definitive or intermediate hosts. In 
all cases there is much to be learned about 
the factors involved in the establishment of 
all associates on and in the shell and with 
the crab itself. Just as the hermit crab is 
attracted to the physical nature of snail 
shells, so too is it likely that the associates 
may show preferences for attachment to 
Shells with a particular topography or 
chemical composition. Likewise, predators 
are attracted by some means (e.g., chemical 
attraction) to certain prey species that pre- 
cede them in their attachment to hermit crab 
shells. 


Acknowledgements 


I am grateful to the following individuals 
from Franklin and Marshall College for 
their help during these studies: K. Miller 
and J. J. Templeton (statistical assistance); 
former student J. P. Studdiford (field and 
laboratory assistance). I am grateful to J. D. 


636 


Williams, University of Rhode Island, who 
kindly read a draft of this paper, D. R. 
Franz, Brooklyn College, New York, for 
clarification of a taxonomic inquiry, and 
Franklin and Marshall College for financial 
support. 


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114(3):640—648. 2001. 


A new species of Exosphaeroma Stebbing (Crustacea: Isopoda: 
Sphaeromatidae) from the Pacific coast of Mexico 


Ma. del Carmen Espinosa-Pérez and Michel E. Hendrickx 


Unidad Académica Mazatlan, ICML, UNAM, PO. Box 811, Mazatlan, Sinaloa 82000, México. 


Abstract.—Exosphaeroma bruscai is described from the Pacific coast of 
Mexico. Exosphaeroma bruscai is most closely related to E. diminuta Menzies 
& Frankenberg, 1966 from the western Atlantic, from which it differs in having 
much longer antenna, an acute rostrum, a triangular first pleopod endopod, a 
transverse suture on exopod of pleopods 3—5, and a considerably longer and 
more slender appendix masculina. Specimens examined here define the geo- 
graphic distribution of E. bruscai from the central Gulf of California to Los 
Arcos, in the southern part of Banderas Bay, Jalisco. Previously published 
records of an unidentified Exosphaeroma from the eastern tropical Pacific in- 
dicate that E. bruscai occurs from sandy beaches of the upper Gulf of Cali- 


fornia to Colombia. 


Sphaeromatidae is the most specious 
family of marine isopods. According to a 
recent survey it contains 633 species (Ken- 
sley & Schotte 2000) and they are often 
very abundant in intertidal and shallow wa- 
ter habitats. Their taxonomy is one of the 
most confused among isopods, in great part 
due to marked sexual dimorphism in some 
genera (Schultz 1969, Brusca 1980, Ken- 
Sley & Schotte 1989) and to the difficulty 
in establishing generic relationships within 
the family (see Bruce 1995). The family has 
been formally divided into five subfamilies 
by Iverson (1982), who based his study on 
previous works by Hurley & Jansen (1977) 
and Bowman (1981). More recently, genera 
of Sphaeromatidae have been reviewed by 
Harrison & Ellis (1991) who presented an 
identification key to genera. 

The genus Exosphaeroma is known from 
the Pacific coast of Mexico, but no identi- 
fied species has been recorded. The first 
published record of Exosphaeroma for the 
area is by Dexter (1972) who recorded E. 
diminuta Menzies & Frankenberg, 1966, a 
west Atlantic sphaeromatid, from sandy 
beaches of the west coast of Panama. Dex- 
ter (1974, 1979) later reported the same 


species for similar habitats in Costa Rica 
and Colombia. This identification, however, 
was considered doubtful by Brusca & Iver- 
son (1985:26—28) who thought that Pacific 
records of Dexter (1974, 1979) belong to 
an undescribed species, probably extending 
from the upper Gulf of California, Mexico, 
to Parque Nacional Santa Rosa, Costa Rica. 
Brusca & Iverson (1985:26—28) provided 
some diagnostic characters of this undes- 
cribed Exosphaeroma, illustrated parts of 
an adult male, but did not give it a new 
name. 

Since Brusca and Iverson’s 1985 note on 
this genus, another report of an Exosphae- 
roma sp. was published by Rios & Ramos 
(1990) from specimens collected in Malaga 
Bay, Colombia. These authors refer to the 
data presented by Brusca & Iverson (1985). 
No further reports on Exosphaeroma from 
the Pacific coast of America have been pub- 
lished. Recent collecting along the Pacific 
coast of Mexico, including sampling on 
sandy beaches, led to the discovery of an 
undescribed species similar to those of 
Brusca & Iverson (1985) in several locali- 
ties. The purpose of the present paper is to 
formalize the description of this apparently 


VOLUME 114, NUMBER 3 


abundant species, the first of the genus Ex- 
osphaeroma reported for the Pacific coast 
of America. 

Abbreviations used in this paper are: St., 
sampling station; TL, total length; NS, un- 
sexed specimen; coll., collector; EMU, Un- 
idad Académica Mazatlan UNAM Inverte- 
brates Reference Collection. 


Sphaeromatidae H. Milne-Edwards, 1840 
Exosphaeroma Stebbing, 1900 
Exosphaeroma bruscai, new species 
Figs. 1—5 


Exosphaeroma diminutum.—Dexter, 1972: 
425; 1974:54; 1979:547 (Pacific records 
only). 

Exosphaeroma sp.—Brusca & Iverson, 
1985:26—28, fig. 1|Ob—d.—Rios & Ramos, 
1990:86-87, fig. 2. 


Type material.—Holotype, 1 male (TL 4.6 
mm), Los Arcos (20°32.5'N, 105°18.4'’W), 
Jalisco, Mexico, 12 Apr 1996 (EMU-4745). 
Paratypes, 3 females (TL 1.9—2.8 mm), Los 
Arcos (20°32.5’N, 105°18.4'’W), Jalisco, 
Mexico, 12 Apr 1996 (EMU-5357). 

Additional material.—El| Tesoro (24° 
18.0’N, 110°19.0’W), La Paz, Southern 
Baja California, Mexico, 17 Jul 1996, 2 
ovigerous females (TL 1.6—2.6 mm) and 1 
unsexed, unmeasured specimen (EMU- 
AVTAS) ee niecdcas Pintas:) Cio: N. i: 
05.5'W), Sonora, Mexico, 24 Mar 1997, 1 
female (TL 4.7 mm) (EMU-4742). Piedras 
Pintas (27°56.5'N, 111°05.5’W), Sonora, 
Mexico, 24 Mar 1997, 2 females (TL 2.9— 
3.4 mm) and 1 ovigerous female (TL 2.4 
mm) (EMU-4744). Bacochibampo Bay 
(27°54.3'N, 110°57.6’W), Guaymas, Sono- 
ra, Mexico, 26 Mar 1997, 1 ovigerous fe- 
male (TL 2.4 mm) collected by diving 
(EMU-5376). 

Description of male.—Body ovate (Fig. 
1A, C), about twice as long as wide; dorsal 
surface smooth. Head wide, rostrum short, 
acute. Length of holotype 4.6 mm. Pereon- 
ite 3 longest, pereonites 1-2 and 4—5 sub- 
equal in length; pereonite 7 more than half 
as long as pereonite 6. Coxae smooth (Fig. 


641 


2), not ventrally directed, without suture, 
not narrowed. Pleon with 3 incompletely 
fused pleonites. Pleotelson smooth, wider 
than long, regularly curving towards pos- 
terior margin in lateral view (Fig. 1B, D); 
wider anteriorly, posteriorly rounded in dor- 
sal view. Epistome scarcely visible in dorsal 
view; elongate in ventral view (Fig. 1B), 
about twice as long as wide in its middle 
part, apex truncate, posterior arms shorter 
than anterior portion. Antennular peduncle 
with 3 articles; flagellum with 8—9 articles. 
Antennal peduncle with 4 articles; flagel- 
lum with 11—12 articles, twice as long as 
antennular flagellum (Fig. 1E). Mandibular 
palp of 3 articles; 2 distal articles with 9 
and 14 spines. Left mandible with incisor 
4-dentate and lacinia mobilis (3-dentate); 
setal row of 4 serrate setae; molar process 
serrate. Right mandible similar in shape, 
with a 4-dentate incisor; setal row of 4 ser- 
rate setae; molar process with dentate mar- 
gin and a seemingly rugose surface (Fig. 
3A, B). Apex of the lateral lobe of maxil- 
lula (Fig. 3C) with 6 serrate setae, and 3 
slender non-plumose, non-serrate setae; me- 
sial lobe with 4 plumose setae, and a short 
non-plumose seta. Maxilla (Fig. 3D) lateral 
lobe with 6 serrate setae, middle with 7, and 
mesial lobe with 7 plumose setae and a 
much longer, slender, mesial plumose setae. 
Maxilliped palp (Fig. 3E) with 4 articles; 
11-10-11-8 setae on articles 1—4, respec- 
tively; endite with 1 retinaculum, 9 plumose 
setae, 4 stout non-plumose setae and about 
7 setules. 

All pereopods with setules, simple 
spines, composed spines (stout spines with 
basal part tipped with a slender seta), and 
diminutive scales on margins. Pereopod 1 
(Fig. 4A) merus slightly shorter than half 
ischium length; ischium with 4 setae at su- 
perior distal angle; merus with 3 setae at 
superior distal angle; carpus triangular, 
short, slightly longer than half merus 
length; propodus shorter than ischium; dac- 
tylus about half propodus length, with | 
subterminal seta on the lower margin; com- 
posed spines, scales and serrate scales pre- 


642 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Exosphaeroma bruscai, new species. A, Holotype, male, dorsal view (EMU-4745); B, pleotelson, 
male, lateral view; C, Paratype, female, dorsal view (EMU-5357); D, pleotelson, female, lateral view; E, ceph- 
alon, male, ventral view; E uropods, dorsal view, detail; G, penes, male. 


VOLUME 114, NUMBER 3 


643 


Fig. 2. Exosphaeroma bruscai, new species, male holotype (EMU-4745). Schematic drawing of body in 


lateral view; appendages not ilustrated. 


sent on articles as illustrated. Pereopods 2— 
3 similar, 3rd slightly longer than 2nd. Pe- 
reopod 3 (Fig. 4B) merus slightly longer 
than half ischium length; ischium with 5 
spines near superior distal angle and 1 
shorter proximal spine; merus with 6 spines 
at superior distal angle; carpus subcylindri- 
cal, similar in length to merus; propodus 
shorter than ischium; dactylus about half 
length of propodus, with 1 subterminal 
spine on the lower margin; composed 
spines, spines, scales and serrate scales pre- 
sent on articles as illustrated. Pereopod 7 
(Fig. 4C) longer and more slender than pe- 
reopods 1—3; merus with 4 spines at supe- 
rior distal angle; carpus distal margin with 
2 inferior and a row of 5 superior serrated 
spines; propodus with 1 bifid spine and 1 
subterminal serrate seta at superior angle; 
composed spines, spines and scales present 
on joints as illustrated. Pereopods 5—6 sim- 
ilar in shape and spination to pereopod 7; 
these pereopods slightly increasing in size 
from 5th to 7th. 

Penes not fused, about twice as long as 
basal width (Fig. 1G). 

Pleopods 1—3 (Fig. 5A—C) endopod and 
exopod with long, plumose marginal setae 
on pleopods as follows (endopod and exo- 
pod): pleopod 1, 16 and 26; pleopod 2, 18 
and 27; pleopod 3, 13 and 30; protopod 
with 3 coupling spines. Distal margin of 
pleopod | exopod slightly curved; endopod 
triangular. Appendix masculina of pleopod 
2 elongate, slender, distally minutely serrate 
and curving, tip rounded, overreaching en- 


dopod of pleopod 2 by about half length of 
the latter. Pleopod 3 as illustrated. Exopod 
of pleopods 3—5 with complete transverse 
suture. Pleopod 4 (Fig. 5D) endopod with 
distinct thickened ridges, 3 short plumose 
marginal setae on ventral margin of exopod 
and 1 on endopod; a row of short non-plu- 
mose setae on the inner margin of exopod. 
Pleopod 5 (Fig. 5E) endopod with distinct 
thickened ridges, a row of short, non-plu- 
mose setae on inner margin of exopod and 
6 similar setae near ventral margin of en- 
dopod; exopod with a deep cleft along inner 
margin, and 5 small lobes with scales close 
to inner distal and ventral margins. 

Uropod exopods of similar length, en- 
dopod slightly longer and acute, exopod 
with rounded tip. 

Female.—Body ovate, about twice as 
long as wide. Pleotelson slightly more in- 
flated than in male, with a weak depression 
near posterior margin. All characters, in- 
cluding all pereopods, very similar to male. 

Etymology.—This species is named for 
Richard C. Brusca, from the BIOSPHERE- 
2 Center, Tucson, Arizona, in recognition of 
his contibution to knowledge of the Isopo- 
da, and for his support of our work. 

Habitat.—In the eastern Pacific, previous 
records of Exosphaeroma are usually from 
sandy beaches. On Panamanian beaches, 
the species was collected in substrate com- 
posed of quartz sand, fine sand and calcar- 
eous shell fragments (Dexter 1972). Rios & 
Ramos (1990) recorded specimens from 
sandy beaches of Colombia. Exosphaeroma 


644 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. 


Exosphaeroma bruscai, new species, male holotype (EMU-4745). A, left mandible; B, right mandible; 
C, right maxilulla; D, right maxilla; E, right maxilliped. 


VOLUME 114, NUMBER 3 645 


“Za 


WWI SE 


Fig. 4. Exosphaeroma bruscai, new species, male holotype (EMU-4745). A, right pereopod 1; B, right 
pereopod 3; C, right pereopod 7. 


646 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ed 


Fig. 5. Exosphaeroma bruscai, new species, male holotype (EMU-4745). A, right pleopod 1; B, right pleopod 
2; C, right pleopod 3; D, right pleopod 4; E, right pleopod 5 (ed = endopod; ex = exopod). 


VOLUME 114, NUMBER 3 


has also been registered in gravels, off man- 
grove lagoons in Costa Rica (Brusca & 
Iverson 1985). In the Gulf of California, 
Mexico, Exosphaeroma has been taken un- 
der shell fragments on sandy stretches of 
tidal flats (Brusca & Iverson 1985). Records 
for E. bruscai are from algae on rocky 
beaches from intertidal zone to at least 3 m; 
environmental data available at the time of 
sampling indicate epibenthic temperature 
range from 20.3 to 28.0°C. 

Although it seems reasonable to assume 
that all specimens of Exosphaeroma previ- 
ously reported from the region belong to E. 
bruscai, the habitat of specimens described 
here differs from the typical sandy beach 
habitat reported previously by Dexter 
(1974, 1979) and Brusca & Iverson (1985). 
Furthermore, three species of Exosphaero- 
ma from the Caribbean are known from 
sand (E£. diminuta) or from algae on rocks 
(E. alba Menzies & Glynn, 1968 and E. 
productatelson Menzies & Glynn, 1968), 
but none has been reported from both hab- 
itats (Kensley & Schotte 1989). Although 
we believe that previous reports of Exos- 
phaeroma from Panama, Costa Rica and 
Colombia correspond to E. bruscai, a com- 
parison of specimens from these localities 
to type specimens of E. bruscai is desirable. 

Distribution.—The species is known 
with certainty from Sonora (27°56.5’N, 
111°05.5’W) to Los Arcos (Banderas Bay), 
Jalisco, on the east coast of the Gulf of Cal- 
ifornia, and from La Paz, South Baja Cali- 
fornia. It is also known from the west coast 
of Costa Rica to Colombia. 

Remarks.—The type species of Exos- 
Phaeroma, Sphaeroma gigas Leach, 1818, 
was recently redescribed (Brandt & Wagele 
1989). Exosphaeroma bruscai expresses the 
typical generic characters considered by 
Brandt & Wagele (1989), in particular the 
ridges or folds of pleopods 4 and 5, and the 
two posteriorly directed submedian flat 
lobes on pleonite 1 (see Bruce 1994). Ex- 
osphaeroma bruscai is very similar to E. 
diminuta which occurs in the western At- 
lantic and with which is was originally con- 


647 


fused. The illustration of the dorsal habitus 
provided by Brusca & Iverson (1985: fig. 
10, B) differs somewhat from our speci- 
mens; the Costa Rica specimens have an 
anteriorly inflated pleon, a character not so 
clearly marked in our specimens. This may 
be due to overshading of the original illus- 
tration, as demonstrated by the lateral view 
of the same pleon (Brusca & Iverson, 1985: 
fig. 10 D) provided by these authors, which 
is much more similar to our illustration (see 
Fig. 1). Comparison of original illustrations 
of E. diminuta by Menzies & Frankenberg 
(1966) with the new species show the fol- 
lowing variations: antenna and antennula 
are similar in size in E. diminuta, while the 
antenna of E. bruscai is almost twice as 
long as the antennula; the rostrum is round- 
ed in E. diminuta, acute in E. bruscai; en- 
dopod of first pleopod is triangular in E. 
bruscai, narrowly oval in E. diminuta; a 
transverse suture is present on exopod of 
pereopods 3—5 in E. bruscai, present only 
on pereopod 4 in E. diminuta; appendix 
masculina slender, much longer than pleo- 
pod 2 endopod in E. bruscai, stouter, just 
reaching apex of the endopod in E. dimi- 
nuta; distal 2 articles of maxilliped palp 
slender in E. diminuta. According to Men- 
zies & Frankenberg (1966: fig. 21 B), clyp- 
eus of E. diminuta is almost rectangular but 
it is figured by Kensley & Schotte (1989: 
fig. 100 H) as being widest in middle 
length, as in E. bruscai. 

Exosphaeroma bruscai also differs from 
other American species of Exosphaeroma 
reported by Kensley & Schotte (1989). Ex- 
osphaeroma alba has a notched, and E. yu- 
catanum (Richardson, 1901) a trilobate, 
margin of the posterior pleotelson; the pleo- 
telson of E. antillense Richardson, 1912 
bears two rounded submedial tubercles; £. 
productatelson, features broad lateral patch- 
es of pigment on pleotelson, not observed 
on any specimen of EF. bruscai. 


Acknowledgments 


The authors thank scientists and students 
who provided assistance during field work. 


648 


Graciano Valenzuela drew figure 1. Mer- 
cedes Cordero did the final editing of the 
manuscript. German Ramirez provided sup- 
port with computing software. Part of this 
study was supported by CONABIO (Com- 
ision Nacional para el Uso y Conocimiento 
de la Biodiversidad), Mexico, project H170. 


Literature Cited 


Bowman, T. E. 1981. Thermosphaeroma milleri and T. 
smithi, new sphaeromatid isopod crustaceans 
from hot springs in Chihuahua, Mexico, with a 
review of the genus.—Journal of Crustacean Bi- 
ology 1:105—122. 

Brandt, A., & J. W. Wagele. 1989. Redescriptions of 
Cymodocella tubicauda Pfeffer, 1887 and Ex- 
osphaeroma gigas (Leach, 1818) (Crustacea, Is- 
opoda, Sphaeromatidae).—Antarctic Science 1: 
205-214. 

Bruce, N. L. 1994. Four new genera of marine isopod 
crustaceans (Sphaeromatidae) from eastern and 
southern Australia—Memoirs of the Museum 
of Victoria 54:399—438. 

. 1995. The taxonomy and phylogeny of tube- 
tailed sphaeromatid isopods (Crustacea) with 
descriptions of new species and a new genus 
from southern Australia.—Ophelia 43:127—180. 

Brusca, R. C. 1980. Common intertidal invertebrates 
of the Gulf of California. The University of Ar- 
izona Press, Tucson, 513 pp. 

, & E. W. Iverson. 1985. A guide to the marine 
isopod crustacea of Pacific Costa Rica.—Revis- 
ta de Biologia Tropical 33:1—77. 

Dexter, D. M. 1972. Comparision of the community 
structures in a Pacific and Atlantic Panamanian 
sandy beach.—Bulletin of Marine Science 22: 
449-485. 

. 1974. Sandy-beach fauna of the Pacific and 

Atlantic coast of Costa Rica and Colombia.— 

Revista de Biologia Tropical 22:51—66. 

. 1979. Community structure and seasonal var- 

iation in intertidal Panamanian sandy beach- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


es.—Estuarine and Coastal Marine Science 9: 
543-558. 

Harrison, K., & J. P. Ellis. 1991. The genera of the 
Sphaeromatidae (Crustacea: Isopoda): a key and 
distribution list.—Invertebrate Taxonomy 5: 
915-952. 

Hurley, D. E., & K. P. Jansen. 1977. The marine fauna 
of New Zealand: family Sphaeromatidae (Crus- 
tacea: Isopoda: Flabellifera)—New Zealand 
Oceanographic Institute Memoir 63:1—80. 

Iverson, E. W. 1982. Revision of the isopod family 
Sphaeromatidae (Crustacea: Isopoda: Flabelli- 
fera) I. Subfamily names with diagnoses and 
key.—Journal of Crustacean Biology 2:248— 
254. 

Kensley, B., & M. Schotte. 1989. Guide to the marine 
isopod crustaceans of the Caribbean. Smithson- 
ian Institution Press, Washington, D.C., 308 pp. 

, & . 2000. World list of marine and 
freshwater Crustacea Isopoda. (URL:http:// 
nmnhwww.si.edu/gopher-menus/WorldListof 
MarineandFreshwaterCrustacealsopoda.html) 

Leach, W. E. 1818. Cymothoadees. in F Cuvier ed., 
Dictionnaire des Sciences Naturelles 12:338— 
354. 

Menzies, R. J., & D. Frankenberg. 1966. Handbook on 
the common marine isopod Crustacea of Geor- 
gia. University of Georgia Press, Athens, 93 pp. 

, & P. W. Glynn. 1968. The common marine 
isopod Crustacea of Puerto Rico.—Studies on 
the Fauna of Curagao and other Caribbean Is- 
lands, 27:1—133. 

Richardson, H. 1901. Key to the isopods of the Atlan- 
tic coast of North America, with descriptions of 
new and little known species.—Proceedings of 
the United States National Museum 23:493— 
579. 

. 1912. Marine and terrestrial isopods from Ja- 
maica.—Proceedings of the United States Na- 
tional Museum 42:187—194. 

Rios, R., & G. E. Ramos. 1990. Los isépodos (Crus- 
tacea: Isopoda) de Bahia Malaga, Colombia.— 
Revista de Ciencias, Universidad del Valle 2: 
83-96. 

Schultz, G. A. 1969. How to know the marine isopod 
crustaceans. W.M.C. Brown Company Publish- 
ers, Dubuque, Iowa, 359 pp. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):649—659. 2001. 


A review of Pseudionella Shiino, 1949 (Crustacea: Isopoda: 
Bopyridae), with the description of a new species parasitic on 
Calcinus hermit crabs from Easter Island 


Christopher B. Boyko and Jason D. Williams 


(CBB) Division of Invertebrate Zoology, American Museum of Natural History, Central Park 
West @ 79th St., New York, New York 10024, U.S.A.; 
(JDW) Department of Biological Sciences, University of Rhode Island, Kingston, 
Rhode Island 02881-0816, U.S.A. (Current address: Department of Biology, Hofstra University, 
Hempstead, New York 11549-1140, U.S.A.) 


Abstract.—Two pairs of bopyrids collected on Easter Island, from the hermit 
crab Calcinus imperialis Whitelegge, belong to the genus Pseudionella Shiino 
(=Pseudasymmetrione Adkinson & Heard, new synonymy) and are described 
as P. akuaku n. sp. This is the first record of any bopyrid isopod from Easter 
Island and the first Pseudionella known to parasitize hermit crabs of the genus 
Calcinus. A review of the species of Pseudionella, with keys to males and 
females, is provided and includes a significant range extension for P. deflexa 


Bourdon. 


Bopyrid isopods found in the branchial 
chambers of hermit crabs are restricted to 
the subfamily Pseudioninae. Seven genera 
have been erected exclusively for species of 
hermit crab branchial parasites, including 
Pseudionella Shiino, 1949, with one Pacific 
and one Atlantic taxon, and Pseudasym- 
metrione Adkinson & Heard, 1978, with 
one Atlantic species. Additionally, nine 
species of the heterogeneous (and probably 
paraphyletic) genus Pseudione Kossman, 
1881, are known from hermit crabs; other 
Pseudione spp. occur on a broad range of 
decapod taxa, including carideans, nephro- 
pids, thalassinoids, galatheoids, and lithod- 
ids. 

One of us (CBB) collected a series of 
hermit crabs from Easter Island (Rapa Nui) 
during August 1999. Two of these crabs 
contained mature female bopyrids, each 
with a male attached to the ventral side of 
the pleonites. Provisional identification of 
these specimens indicated that they ap- 
peared to belong to the genus Pseudasym- 
metrione Adkinson & Heard. However, an 
examination of Pseudasymmetrione type 


material showed that genus to be a syno- 
nym of Pseudionella Shiino, and our spec- 
imens cannot be placed into any of the three 
described species now placed in that genus. 
These specimens are described as a new 
species of Pseudionella which is most 
closely related to an Atlantic congener, P. 
markhami (Adkinson & Heard). 


Methods 


Hermit crabs inhabiting gastropod shells 
were collected intertidally on Easter Island 
during August 1999. The data label in the 
vial containing the parasitized specimens 
was badly damaged during shipping of 
specimens from Easter Island to New York 
and, consequently, no specific locality can 
be determined. Specimens were preserved 
in 70% ethanol. The shells were cracked 
using a vise and the crabs removed and ex- 
amined for parasites. Collection of a single 
female Pseudionella deflexa was made on 
Andros Island, Bahamas in September 2000 
using methods identical to those above. 

Drawing tube sketches made of the spec- 


650 


imens were scanned into a Macintosh™ 
computer. Images were then prepared using 
the programs Adobe Photoshop® and Ado- 
be Illustrator®. 

Shield length (SL) is provided as an in- 
dicator of size for the host crabs. Isopod 
size is given as total body length (anterior 
margin of head to posterior margin of pleo- 
telson). Measurements were made to 0.01 
mm using an ocular micrometer. 

Specimens of the new species and P. de- 
flexa are deposited in the Division of Inver- 
tebrate Zoology, American Museum of Nat- 
ural History, New York, U.S.A. (AMNH). 
Specimens of Pseudasymmetrione markhami 
were borrowed from the National Museum 
of Natural History (Naturalis), Leiden, the 
Netherlands (RMNH) and the National Mu- 
seum of Natural History, Smithsonian In- 
stitution, Washington, D.C. (USNM). 


Family Bopyridae Rafinesque, 1815 
Subfamily Pseudioninae Codreanu, 1967 
Genus Pseudionella Shiino, 1949 


Pseudionella Shiino, 1949:60—61 (type 
species by monotypy: Pseudionella at- 
tenuata Shiino, 1949). 

*““Undescribed genus” Markham, 1978:110. 
Pseudasymmetrione Adkinson & Heard, 
1978:409, 412 (type species by monoty- 
py: Pseudasymmetrione markhami Ad- 
kinson & Heard, 1978) (new synonymy). 

not Pseudionella, Shiino, 1958:41 (=Bo- 
pyrissa Nierstrasz & Brender a Brandis, 
LOS): 


Diagnosis.—Female with seven pereon- 
ites and five pleonites plus pleotelson, all 
laterally distinct. Frontal lamina entire. Lat- 
eral plates very short. Marsupium enclosed 
by well-developed oostegites. Five pairs of 
lamellar pleopods, anterior two pairs bira- 
mous with exopodite distally bilobed; pos- 
terior three pairs uniramous. Uropoda uni- 
ramous. Male with seven distinct pereonites 
and five distinct pleonites plus pleotelson, 
pleonites markedly reduced in width from 
pereonites. Five pairs of uniramous tuber- 
culiform or flap-like pleopods. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Remarks.—The addition of the new tax- 
on described below to Pseudionella, and the 
synonymy of Pseudasymmetrione, requires 
some modifications to the original diagnosis 
of Shiino (1949). The frontal lamina is en- 
tire, although Shiino (1949) called it rudi- 
mentary in the type species. The first two 
pairs of pleopods in the female are bira- 
mous with the endopodite bilobed at the tip, 
while the last three pairs are uniramous. 
Both Shiino (1949) and Adkinson & Heard 
(1978) considered the coalescence of the 
head with the first pereonite in the male to 
be of great importance, but this character 
can vary between individuals within a spe- 
cies (e.g., Stegophryxus hyphalus Mark- 
ham, 1972:38) and we are unconvinced in 
its reliability as a diagnostic feature. The 
pleotelson of the male is either a simple 
cone (as in P. attenuata and P. deflexa) or 
has lateral projections (as in P. markhami 
and the new species) which may represent 
reduced lateral plates on the sixth pleonite 
(=pleotelson). 

The synonymy of Pseudasymmetrione 
with Pseudionella was suggested, but not 
formally proposed, by Bourdon (1979). 
Bourdon (1979) noted that the one, perhaps 
significant, difference between the two taxa 
was the form of the male pleotelson (simple 
in Pseudionella and with fused lateral 
plates in Pseudasymmetrione) but won- 
dered if this was sufficient for the estab- 
lishment of a separate genus. We agree that 
the four characters given as diagnostic for 
Pseudasymmetrione by Adkinson & Heard 
(1978) do not appear to warrant separation 
of the genera: the female dorsal segmenta- 
tion is medially indistinct in both P. mark- 
hami and the new species, but is variable 
between the specimens of the new species, 
and therefore not a good character; the fron- 
tal lamina of Pseudionella is not reduced or 
absent, as stated by Shiino (1949), and there 
is little difference in this character between 
the four species; the coxal plates of P. 
markhami are well developed, but are re- 
duced in the other taxa including the new 
species; and contrary to the statement of 


VOLUME 114, NUMBER 3 


Adkinson & Heard (1978), males of P. 
markhami do possess pleopods, but of a 
form more reduced than those seen in P. 
attenuata and P. deflexa. The only character 
left to discriminate between the two genera 
is the shape of the male pleotelson which, 
as noted by Bourdon (1979), was not in- 
cluded in the original diagnosis of Pseu- 
dasymmetrione. In our opinion, this single 
difference does not warrant separate generic 
status for these species. We propose that 
Pseudionella contains two species-groups, 
each with one Atlantic and one Pacific tax- 
on: P. attenuata and P. deflexa; P. mark- 
hami and the new species from Easter Is- 
land. Note that Pseudionella pyriforma Shi- 
ino, 1958, does not belong in this genus, 
but to Bopyrissa Nierstrasz & Brender a 
Brandis, 1931 (see Bourdon 1979, Mark- 
ham 1982). Pseudionella appears closely 
allied to the diverse genus Pseudione which 
contains species with very similar female 
morphology, but with five pairs of biramous 
pleopods in the females (e.g., Nierstrasz & 
Brender a Brandis 1931, Shiino 1933). The 
males of Pseudionella also resemble those 
of Pseudione, but have more laterally con- 
stricted pleonites. Although Pseudionella 
contains several clearly apomorphic char- 
acters (€.g., uniramous pleopods 3-5 in fe- 
males, narrowed pleonites in males) as 
compared to Pseudione, a detailed phylo- 
genetic analysis is needed to fully under- 
stand the relationships between these gen- 
era and the others in the subfamily. 


Pseudionella attenuata Shiino, 1949 


Pseudionella attenuata Shiino, 1949:62— 


63, fig. 2. 


Material examined.—unavailable. 

Type locality.—Seto, Wakayama Prefec- 
ture, Japan (Shiino 1949, type specimens (if 
extant) in National Science Museum, To- 
kyo). 

Distribution and host.—Japan, on Pa- 
gurus sp. (Shiino 1949, as Eupagurus sp.). 

Remarks.—No specimens have been re- 
ported subsequent to the two type speci- 


651 


mens. Unfortunately it has not been possi- 
ble to directly examine the types, nor to 
even confirm if they are still extant, as Shi- 
ino’s material is in a very poor state of or- 
ganization (M. Takeda, pers. commun. 15 
Jun 1999). The female type was dextral, 
which suggests that it came from the right 
branchial chamber of the host. 


Pseudionella markhami (Adkinson & 
Heard, 1978), new combination 


““Undescribed ... 
110. 

Pseudasymmetrione markhami Adkinson & 
Heard, 1978:412—414, figs. 1—-3.—Mark- 
ham, 1988:8-9, fig. 2. 

*““Pseudasymmetrione ... undescribed spe- 
cies” Adkinson & Heard, 1978:417. 


species’”” Markham, 1978: 


Material examined.—Holotype: sinistral 
female (3.14 mm) infesting left branchial 
chamber of Pagurus annulipes (Stimpson) 
(sex and size unknown), Morehead Chan- 
nel, Carteret County, North Carolina, 
U.S.A., R. W. Heard coll., 26 Aug 1963 
(USNM 170590). ““Allotype’’: male (0.99 
mm) same data as holotype (USNM 
170590). Paratypes: female (2.14 mm) and 
male (0.89 mm) infesting male host (1.7 
mm SL), female (2.00 mm) and male (0.78 
mm) infesting male host (1.7 mm SL), fe- 
male (2.86 mm) and male (0.87 mm) in- 
festing female host (2.1 mm SL), all fe- 
males sinistral and infesting left branchial 
chambers of P. annulipes, Morehead Chan- 
nel, Carteret County, North Carolina, 
U.S.A., R. W. Heard coll., 25 Jun 1970 
(USNM 170593); non-types: dextral female 
(3.14 mm), and male (0.99 mm), infesting 
[presumed right] branchial chamber of /ri- 
dopagurus iris (A. Milne Edwards) (sex 
and size unknown), Sta. P-757, 11°41'N, 
69°21'W, off Peninsula de Paraguana, Ven- 
ezuela, 161—187 m, R/V Pillsbury coll., 27 
Jul 1968 (USNM 172224); sinistral female 
(3.14 mm), male (1.57), sinistral female 
(1.26 mm), male (0.72 mm) infesting Pa- 
gurus brevidactylus (Stimpson) (host sexes, 
sizes and infested branchial chambers un- 


652 


known), Punta de Betin, Santa Marta, Dept. 
Magdalena, Colombia, under stones, 1—2 m, 
coll. H. G. Miiller, 4 Feb 1986 (RMNH 
7042); sinistral female (1.26 mm), infesting 
Pagurus stimpsoni (A. Milne Edwards & 
Bouvier) (host sex, size, and infested bran- 
chial chamber unknown), Punta de la 
Aquia, ca. 4 km east of Santa Marta, Dept. 
Magdalena, Colombia, on coral rubble, 17— 
19 m, coll. H. G. Miiller, 9 Jan 1986 
(RMNH 7058). 

Type locality.—Morehead Channel, Car- 
teret County, North Carolina, United States 
(Adkinson & Heard 1978, type specimens 
in USNM, Zoologiske Museum Copenha- 
gen, and the collection of D. Adkinson). 

Distribution and hosts.—North Carolina, 
U.S.A., on Pagurus annulipes (Stimpson) 
(Adkinson & Heard 1978, herein); Vene- 
zuela, on Iridopagurus iris (A. Milne Ed- 
wards) (Markham 1978, Adkinson & Heard 
1978, herein); Colombia (Atlantic), on Pa- 
gurus brevidactylus (Stimpson) and P. 
stimpsoni (A. Milne Edwards & Bouvier) 
(Markham 1988, herein). 

Remarks.—The pereonites and pleonites 
of male P. markhami are much more com- 
pacted than in the other three taxa in the 
genus, but the shape of the male pleotelson 
and the form of the male pleopods are very 
close to the new species from Easter Island. 
Although Adkinson & Heard (1978) did not 
designate an allotype, the male accompa- 
nying the female holotype is of equivalent 
status. Markham (1988) correctly identified 
the above cited specimens of Pseudionella 
markhami from Colombia, but inaccurately 
drew the male pleotelson of RMNH 7042 
as having two slender projections; the shape 
is identical to that of the male allotype and 
of the other specimens in RMNH. Adkinson 
& Heard (1978) indicated that Markham’s 
(1978:110) “undescribed new genus and 
species”’ from Venezuela appeared to be an 
undescribed Pseudasymmetrione (=Pseu- 
dionella). However, examination of those 
two specimens indicated no notable differ- 
ences from P. markhami, other than the 
unique dextral orientation of the female. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Pseudionella deflexa Bourdon, 1979 


Pseudionella deflexa Bourdon, 1979:139— 
141, fig. 1. 


Material examined.—sinistral female 
(2.5 mm, with one female and three male 
Cabirops cryptoniscid isopods in the brood 
chamber), infesting male Pagurus brevidac- 
tylus (Stimpson) (2.1 mm SL), Station 
AO9R, 24°53'13.3"N, 77°54'47.2"W, sand/ 
algal plain, 1.2—1.8 m, Andros Island, Ba- 
hamas, coll. C. B. Boyko, 1 Sep 2000 
(AMNH 18204). 

Type locality.—24°35'5"S, 46°31'W, 45 
m, Brazil (Bourdon 1979). 

Distribution and hosts.—Brazil, on Pa- 
gurus criniticornis (Dana) (Bourdon 1979); 
Bahamas, on Pagurus  brevidactylus 
(Stimpson) (herein). 

Remarks.—The Bahamas specimen rep- 
resents a new host record and a significant 
range extension for the species, which was 
previously known only from Brazil. The fe- 
males of this species are extremely close to 
those of P. markhami, but can be distin- 
guished by more delineated segmentation of 
the pereonites, less pronounced coxal plates 
(especially on the posterior pereonites), and 
less pronounced lateral projections on the 
pleotelson. The males of the two species are 
easily distinguished by the presence (P. 
markhami) or absence (P. deflexa) of lateral 
projections on the pleotelson and by the dif- 
ferent forms of the pleopods. 


Pseudionella akuaku, new species 
Figs. 1—4 


Material examined.—Holotype: sinistral 
female (1.44 mm), infesting left branchial 
chamber of male Calcinus imperialis Whi- 
telegge (2.0 mm SL; AMNH 18187), in- 
habiting shell of Planaxis akuana Rehder, 
Easter Island, intertidal, coll. C. B. Boyko, 
Aug 1999 (AMNH 18201). Allotype: male 
(0.91 mm), same data as holotype (AMNH 
18202). Paratypes: sinistral female (1.75 
mm), male (1.26 mm) infesting left bran- 
chial chamber of male Calcinus imperialis 


VOLUME 114, NUMBER 3 


(2.0 mm SL; AMNH 18187), inhabiting 
shell of Planaxis akuana Rehder, Easter Is- 
land, intertidal, coll. C. B. Boyko, Aug 
1999 (AMNH 18203). 

Type locality.—Easter Island, territory of 
Chile, Pacific Ocean. 

Description.—Female (Figs. 1, 2), based 
on holotype: Body length 1.44 mm, maxi- 
mal width 1.19 mm, head length 0.48 mm, 
head width 0.52 mm, pleon length 0.45 
mm. Pereon weakly S-shaped; head deflect- 
ed to the right and pleon weakly to the left. 
Body outline broad at pereon, narrow at 
pleon and elongated (Figs. 1A, B). Dark 
spot or band of pigmentation at junction of 
all coxal plates and pereonites (in dorsal 
view). 

Head weakly produced with anterior lam- 
ina raised and recurved along distal margin. 
Eyes present, large relative to cephalon, oc- 
curring near posterolateral corners of raised 
lamina edge (but absent in paratype fe- 
male). Antenna (Fig. 2A) of three articles; 
antennule (Fig. 2A) of three articles, all 
segments with fine scales bearing setae on 
distal margins, distal margins of segments 
with setae. Maxilliped (Fig. 2B) with elon- 
gate rounded spur; palp subcircular with 
distal narrow, rounded projection; posterior 
margin setose. First oostegite proximal lobe 
ovate, distal lobe subtriangular, internal 
ridge smooth (Figs. 2C, D). 

Pereon broadest across pereonite 4, ta- 
pering anteriorly and posteriorly. Coxal 
plates on sides of pereonites all similar but 
larger on left side of body. Oostegites 
sparsely covered in minute tubercles; pos- 
teriormost oostegite with fringe of setae on 
posterior margin. Pereopods 3—6 of about 
Same size; pereopods 1, 2, and 7 (Fig. 2E, 
F) slightly smaller and shorter. Dorsal mar- 
gins of dactylus and basis, and ventral mar- 
gins of propodus, carpus, and merus with 
numerous plate-like scales bearing setae on 
distal margins; short setae at distal tip of 
propodus. First two pereopods surrounding 
head region; no large gaps between any pe- 
reopods. 

Pleonites | and 2 with extended lanceo- 


653 


late, distally rounded, biramous pleopods 
and uniramous short lateral plates; exopod- 
ite of pleopod 1 bilobed, endopodite smaller 
and entire; exopodite and endopodite of 
pleopod 2 entire; pleonites 3—5 with lan- 
ceolate, uniramous pleopods and uniramous 
short lateral plates (Fig. 2G); lateral plates 
slightly reduced and pleopods markedly re- 
duced in size from anterior to posterior; 
pleotelson (Fig. 2H) appears trifid distally 
as a result of incomplete fusion of segment 
with sixth pair of lateral plates, all projec- 
tions subequal in length and width, with 
pair of large broad lanceolate, distally 
rounded, uropods. Lateral plates and uro- 
pods with dense covering of scales bearing 
setae on distal margins (Fig. 21). 

Male (Figs. 3, 4), based on allotype: 
Length 0.91 mm, head length 0.13 mm, 
head width 0.28 mm, pleon length 0.21 
mm. Occurring on ventral side of pleon of 
female; directed anteroposteriorly. 

Head suboval, widest posteriorly, distinct 
from with first pereonite (Figs 3A, B). 
Large eyes (relative to cephalon) near pos- 
terolateral margin. Antenna of five articles, 
distally setose; extending posterolaterally 
from head; antennule of three articles; an- 
tennae and antennule with scattered small 
scales bearing setae on distal margins (Fig. 
4A). 

Pereonites 3 and 4 broadest, tapering an- 
teriorly and posteriorly. All pereonites di- 
rected laterally. Irregular dark pigmentation 
pattern at junction of body and pereonite 
lateral projections and on first three pleon- 
ites. All pereopods (Figs. 4B, C) subequal, 
all articles distinctly separated, no scales 
apparent on dorsal or ventral surfaces. 

Pleonites tapering posteriorly and direct- 
ed laterally. All pleonites distinctly seg- 
mented, weakly produced laterally and 
markedly narrower than pereonites. No 
midventral tubercles (Figs. 3B, 4D). Pleo- 
telson (Fig. 4D) notched medially with mi- 
nute anal cone, produced distolaterally into 
rounded lobes, distolateral corners of lobes 
with scales and setae; uropods absent. 

Distribution and host.—On hermit crabs, 


654 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 
ventral view. Scale = 500 pm. 


Pseudionella akuaku, new species. Female 


Calcinus imperialis from Easter Island; in- 
tertidal. 

Etymology.—The specific name akuaku 
is derived from the Rapa Nui word for su- 
pernatural beings who sometimes assumed 
animal shapes (Blixen 1993). The name is 
used an a noun in apposition. 

Remarks.—The pleopodal formula of the 
female, the attenuated width of the pleoni- 
tes of the male as compared with the per- 
eonites, and the presence of uniramous ple- 
opods in the male, clearly place this taxon 
in Pseudionella. Additionally, females of 
all species of Pseudionella possess small 


, 1.44 mm, AMNH 18201, holotype. A, dorsal view; B, 


lobes on the ventral surface of each pereon- 
ite mesial to the pleopods. The nature of 
these lobes is unknown but, as pointed out 
by Shiino (1949), they cannot be reduced 
pleopods as they occur alongside fully bi- 
ramous pleopods. A comparison with the 
other three species now placed in this genus 
shows several important differences be- 
tween the taxa. Pseudionella akuaku n. sp. 
has a “‘trilobed’’ appearance in the pleotel- 
son of the female (i.e., a partly fused sixth 
pair of lateral plates), very broad uropods 
of the female, pronounced lateral lobes 
(fused sixth lateral plates) on the pleotelson 


VOLUME 114, NUMBER 3 655 


Fig. 2. Pseudionella akuaku, new species. Female, 1.75 mm, AMNH 18203, paratype (A—F); Female, 1.44 
mm, AMNH 18201, holotype (G—I). A, left antenna and antennule; B, right maxilliped, external; C, right 
oostegite 1, external; D, right oostegite 1, internal; E, right pereopod 1; E right pereopod 7; G, posterolateral 
view, left side; H, pleotelson and uropods; I, uropod detail (EN = endopod; EX = exopod; L = lateral plate; 
P = pleopod; PT = pleotelson; U = uropod; numbers indicate pleonite). Scale = 150 wm (1), 200 pm (A, E, 
F), 225 wm (H), 250 pm (G), 300 pm (B), and 500 wm (C, D). 


656 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. 
ventral view. Scale = 250 pm. 


of the male, and flap-like pleopods. In con- 
trast, both P. attenuata and P. deflexa have 
slender uropods on the female, no lateral 
lobes on the pleotelson of the male, and 
tuberculiform pleopods. The pleotelson of 
P. deflexa does have a “‘trilobed”’ appear- 
ance, but it is less pronounced than that of 
P. akuaku n. sp. Pseudionella markhami 
possesses both flap-like pleopods on the 
male and broad uropods on the female, 
while the male has only weakly produced 
lateral lobes on the pleotelson. The anten- 
nae and antennules of female P. deflexa, P. 
markhami, and P. akuaku n. sp. are each 
composed of three articles, while those of 
P. attenuata are 2 and 3 segmented, re- 
spectively (Shiino 1949). 

Female and male specimens of Pseudi- 
onella spp. possess plate-like scales bearing 


Pseudionella akuaku, new species. Male, 0.91 mm, AMNH 18202, allotype. A, dorsal view; B, 


setae on the distal margins of the antennae, 
antennules, and pereopods. Such scales 
have been noted in other bopyrids and their 
fine structure has been examined by SEM 
in Heterocepon marginatum Shiino, 1936, 
by Janssen & Brandt (1994), who suggested 
that these scales may aid in attachment to 
host crabs (for females) or female bopyrids 
(for males). 

The shape of the female pleotelson in all 
of the nine species of Pseudione found on 
hermit crabs is nondescript, typically being 
a small rounded or slightly pointed lobe, as 
is true of Pseudionella attenuata. In marked 
contrast, the female pleotelsons of Pseudi- 
onella deflexa, P. markhami, and P. akuaku 
n. sp. are large and trifid with pronounced 
and rounded median and lateral lobes. 
These lateral lobes represent poorly devel- 


VOLUME 114, NUMBER 3 


657 


Fig. 4. Pseudionella akuaku, new species. Male, 1.26 mm, AMNH 18203, paratype (A); Male, 0.91 mm, 
AMNH 18202, allotype (B—D). A, right antenna and antennule; B, left pereopod 1; C, right pereopod 7; D, 
ventral view of posterior pleonites and pleotelson. Scale = 50 wm (B, C), 100 pm (A, D). 


oped lateral plates partly fused with the 
pleotelson and are quite different from the 
lateral plates found on pleonites 1—5. Pseu- 
dionella attenuata may represent a transi- 
tional form between the hermit crab-infest- 
ing Pseudione and the other species of 
Pseudionella. 

Ecology.—A total of 81 hermit crabs 
were collected from intertidal and subtidal 
locations on Easter Island from 22—31 Au- 
gust 1999. Examples of both previously 
known shallow water hermit crabs from the 
island were present in the collection: Cal- 
cinus pascuensis Haig (18 specimens) and 
C. imperialis (65 specimens), as well as a 
newly recorded species, C. vachoni Forest 
(1 specimen). Only two male hermit crab 


specimens of C. imperialis were found to 
have bopyrid parasites (2.4% overall prev- 
alence). This is the first report of a member 
of Pseudionella on a species of Calcinus, 
the other three species being known from 
Pagurus spp. (Shiino 1949, Adkinson & 
Heard 1978, Bourdon 1979). 


Key to females of the species of 
Pseudionella 


ll, Pleoiellsom SiS .5esse05548 P. attenuata 
— Pleotelson “‘trifid’’ (with lateral plates) 
2. Medial pereonite segmentation entire 

ns \ofosecacigchecir dle etechicat isa eg oe ae P. deflexa 
— Medial pereonite segmentation indistinct 


658 


3. Pleotelson lateral plates subacute, ex- 
tending well beyond apex of pleotelson 
eNO iis LOR RE I eae P. markhami 


— Pleotelson lateral plates rounded, sube- 
qual to apex of pleotelson ......... 


Key to males of the species of 
Pseudionella 


1. Pleotelson with lateral projections .... 2 
— Pleotelson simple, without lateral projec- 
tions 
2. Pleotelson lateral projections large, dis- 
tinct P. akuaku n. sp. 
— Pleotelson lateral projections small, in- 


GISHING tet eens a ton as P. markhami 
3. Pleotelson minute, shorter than pleonite 

Meee sore felts cos nv tieree etree he P. deflexa 
— Pleotelson elongate, longer than pleonite 

Oa ea ae Re, Seseieny a tty eae P. attenuata 


Acknowledgments 


Joseph Poupin (Institut de Recherche de 
1’ Ecole Navale, France) confirmed the iden- 
tifications of two Calcinus spp. and identi- 
fied C. vachoni. C. H. J. M. Fransen 
(RMNH) and Rafael Lemaitre (USNM) 
kindly provided specimens on loan. Fund- 
ing for the Invertebrate Survey of Easter 
Island was provided to CBB by the U. S. 
National Park Service (USNPS), Gateway 
National Recreation Area, Division of Nat- 
ural Resources, as part of a Science Muse- 
um of Long Island/Explorers Club five-year 
research expedition to explore the impacts 
of El Nifio events on World Heritage Sites, 
through the efforts of John Tanacredi 
(USNPS). The Bahamas material was ob- 
tained from work supported by NASA un- 
der award no. NAG5-8543 and by the Cen- 
ter for Biodiversity and Conservation at the 
American Museum of Natural History, 
through the efforts of Daniel Brumbaugh 
(AMNH) and Paula Mikkelsen (AMNH). 
Two anonymous reviewers contributed 
greatly to the final version of this publica- 
tion. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Literature Cited 


Adkinson, D. L., & R. W. Heard. 1978. Description of 
a new genus and species of Pseudioninae (Iso- 
poda: Bopyridae) parasite of the hermit crab 
Pagurus annulipes (Stimpson) from North Car- 
olina.—Proceedings of the Biological Society 
of Washington 91:408—417. 

Blixen, O. 1993. The concept of akuaku. Pp. 138-142 
in S. R. Fischer, ed., Easter Island Studies. Con- 
tributions to the History of Rapanui in Memory 
of William T: Mulloy. Oxbow Monograph 32. 

Bourdon, R. 1979. Crustacés isopodes: Bopyridae par- 
sites de pagures.—Annales de I’ Institut Océan- 
ographie, n.s. 55 (suppl.):139—144. 

Codreanu, R. 1967. Classificarea evolutiva a bopirien- 
ilor, isopode parazite ale crustaceelor decapode 
si importanta lor biologica generala.—Studii si 
Cercetari de Biologie 19:203—211. 

Janssen, H., H., & A. Brandt. 1994. Heterocepon mar- 
ginatum Shiino, 1936, (Crustacea: Isopoda: Ep- 
icaridea: Bopyridae) a new hyperparasite from 
the Philippines, and a short review of the biol- 
ogy of the Bopyridae.—The Philippine Scientist 
31:5-31. 

Kossman, R. 1881. Studien tiber Bopyriden. I, Gigan- 
tione moebii und Allgemeines tiber die Mund- 
werkzeuge der Bopyriden. Il, Bopyrina virbii, 
Beitrage zur Kenntnis der Anatomie und Meta- 
morphose der Bopyriden.—Zeitschrift fiir Wis- 
senschaftliche Zoologie, Leipzig 35:652—680. 

Markham, J. C. 1972. Parasitic bopyrid isopods of the 
amphi-Atlantic genus Stegophyrxus Thompson 
with the description of a new species from Cal- 
ifornia.—Bulletin of the Southern California 
Academy of Sciences 73:33—41. 

1978. Bopyrid isopods parasitizing hermit 

crabs in the northwestern Atlantic Ocean.—Bul- 

letin of Marine Science 28:102—117. 

. 1982. Bopyrid isopods parasitic on decapod 

crustaceans in Hong Kong and southern China. 

Pp. 325-391 in B. S. Morton & C. K. Tseng, 

eds., The Marine Flora and Fauna of Hong 

Kong and Southern China, vol. 1. Hong Kong 

University Press. 

. 1988. Descriptions and revisions of some spe- 
cies of Isopoda Bopyridae of the north western 
Atlantic Ocean.—Zoologische Verhandelingen 
Uitgegeven Door Het Rijksmuseum van Na- 
tuurlijke Historie te Leiden 246:63 pp. 

Nierstrasz, H. KF, & G. A. Brender a Brandis. 1931. 
Papers from Dr. Th. Mortensen’s Pacific Expe- 
dition 1914-16. LVII. Epicaridea IJ—Viden- 
skabelige Meddedelser fra den Dansk Naturhis- 
toriske Forening i K@benhavn 91:147—226, 1 
plate. 

Rafinesque-Schmaltz, C. S. 1815. Analyse de la nature 
ou tableau de l’univers et des corps organisés. 
Palermo, 224 pp. 


VOLUME 114, NUMBER 3 659 


Shiino, S. M. 1933. Bopyrids from Tanabe Bay.— 


. 1949. On two new genera of Bopyridae found 


Memoirs of the College of Science, Kyoto Im- in Japan.—Bulletin of the Biogeographical So- 
perial University Series B 8:249—300. ciety of Japan 14:57—-63. 
. 1936. Bopyrids from Shimoda and other dis- . 1958. Note on the bopyrid fauna of Japan.— 


tricts.—Records of Oceanographic Works in Ja- Report of Faculty of Fisheries, Prefectural Uni- 
pan 8:161—176. versity of Mie 3:29-74, pl. 3. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):660—666. 2001. 


Foliomolgus cucullus, a new genus and species of Clausidiidae 
(Crustacea: Copepoda: Poecilostomatoida) associated with a 
polychaete in Korea 


Ul-Hoi Kim 


Department of Biology, Kangnung National University, Kangnung 210-702, Republic of Korea, 
e-mail: ihkim @kangnung.ac.kr 


Abstract.—Foliomolgus cucullus, new genus and species, is described as an 
associate of the polychaete Marphysa sanguinea (Montagu) inhabiting the in- 
tertidal sands in Jeju Island in Korea Strait. The new genus possesses primitive 
features such as the complete segmentation and setation of rami on legs 1—4, 
7-segmented antennule, 4-segmented antenna, and 5-segmented female uro- 
some. It possesses also characteristic features such as a spatulate distal segment 
of the maxilla, foliaceous ventral elements and rudimentary dorsal claw on the 
mandible, atrophied distal segment of the antenna, and no maxilliped in the 


female. 


The copepods of the family Clausidiidae 
have been considered to be the most ances- 
tral group of the order Poecilostomatoida. 
They are found on/in various tubicolous in- 
vertebrates, including crustaceans, bivalves, 
and polychaetes. 

The host of the new genus, Marphysa 
sanguinea (Montagu), is a cosmopolitan 
species of Polychaeta occurring in warm 
seas (Imajima & Hartman, 1964). Kim 
(2000) reported Clausia lobata as a new spe- 
cies of copepod associated with M. sangui- 
nea from the Yellow Sea. During a recent 
field survey in Jeju Island, the largest island 
in Korea, a number of the same polychaete 
species were collected from the intertidal 
sands, and searched for copepod associates. 
At this island C. lobata was not found on 
the same polychaete, but, instead, a new spe- 
cies of Copepod belonging to a new genus 
of the family Clausidiidae was collected. Al- 
though only one female and two males were 
found, they were large enough to facilitate a 
thorough study, leaving type specimens 
available for future study. 

Before microscopic observation and dis- 
section, copepod specimens were cleared in 


lactic acid. Dissections were done using the 
reversed slide method (Humes & Gooding, 
1964). Drawing was done with the aid of a 
camera lucida. In the description of species, 
the body lengths were measured from the 
anterior tip of the cephalothorax to the pos- 
terior margin of the caudal rami, excluding 
the caudal setae. In the formula for the ar- 
mature of legs 1—4 the Roman numerals in- 
dicate spines and the Arabic numerals rep- 
resent setae. 


Family Clausidiidae Embleton, 1901 
Foliomolgus, new genus 


Diagnosis.—Clausidiidae. Body cyclo- 
piform, relatively large, 9-segmented in fe- 
male and 10-segmented in male. Prosome 
composed of cephalothorax and 3 meta- 
somites. Fifth pedigerous somite of female 
with dorsal hood. Antennule 7-segmented; 
setation as for the species. Antenna 4-seg- 
mented; third segment with | claw and 3 
setae; terminal segment atrophied, armed 
with 4 spiniform setae and 3 simple setae. 
Labrum reduced, incompletely covering 
oral appendages. Mandible with atrophied 
terminal process and 3 foliaceous subter- 


VOLUME 114, NUMBER 3 


minal elements. Paragnath encircling 
mouth, with 2 lobate processes on inner 
margin. Maxillule distally bilobed, with 3 
and 5 setae on respective lobes. Maxilla 2- 
segmented, simplified, and armed with 1 
seta on basal segment and 2 setae on digi- 
tiform distal segment. Maxilliped absent in 
female. Male maxilliped of well-developed 
and composed of 4 segments, including ter- 
minal claw. Legs 1—4 with 3-segmented 
rami; setation as for the species. Leg 5 with 
1 seta on basal segment and 4 setae on dis- 
tal segment. Basal segment of male leg 5 
fused with fifth pedigerous somite. Male leg 
6 represented by | seta on genital flap. 

Etymology.—The generic name Foliom- 
olgus is a combination of folium (=leaf in 
Latin) and molgus (the ending of many ge- 
neric names of copepod associates). It al- 
ludes to the foliaceous subterminal ele- 
ments on the mandible in both sexes. The 
gender is masculine. 

Type species.—Foliomolgus cucullus, 
new species 


Foliomolgus cucullus, new species 
Figs. 1-3 


Type specimens.—One female and two 
males found on the external surface of the 
polychaete Marphysa sanguinea (Montagu) 
collected from the intertidal coral sands at 
Sinhung-ri on the northern shore of Jeju Is- 
land (approximately 33°33’N, 126°39’E), on 
3 Jun 2000. Holotype female (USNM 
309082; left antennule, left antenna, and oral 
appendages dissected out and mounted on a 
slide) and allotype (USNM 309083; an intact 
male) have been deposited in the National 
Museum of Natural History, Smithsonian In- 
stitution. Dissected paratype (one male) is re- 
tained in the collection of the author. 

Female.—Body (Fig. 1A) cyclopiform, 
2.67 mm long. Greatest width 1.02 mm. 
Prosome composed of cephalothorax and 
three pedigerous somites. Rostral area of 
cephalosome produced anteriorly. Prosomal 
somites with well developed epimera. Epi- 
mera of second and third pedigerous so- 


661 


mites with pointed posterior corners. Epi- 
meron of fourth pedigerous somite with 
rounded corners. Urosome (Fig. 1C) slender 
and five-segmented. Fifth pedigerous so- 
mite 405 jm wide, characteristically with 
dorsal hood (487 wm wide) covering most 
of dorsal surface of fifth pedigerous somite 
and anterior part of genital double-somite 
(Fig. 1B). Genital double-somite 335 < 295 
lum, widest in anterior third, then tapering 
posteriorly. Genital area located laterally, 
invisible from dorsal and ventral views. 
Three abdominal somites 210 X 165, 162 
x 142, and 115 X 115 pm, respectively. 
Anal somite with spinules along posterov- 
entral border. Caudal ramus slender, diver- 
gent, 221 X 40 pm (5.53:1), with parallel 
lateral margins and 6 setae. Posteroventral 
border of caudal ramus armed with several 
denticles (Fig. 1D). All caudal setae naked. 
One of terminal setae distinctly larger than 
others, 1.05 mm long. 

Egg sac not seen. 

Rostrum wider than long, rounded pos- 
terior margin (Fig. 1E). Antennule (Fig. 1F) 
7-segmented, 731 wm long, with armature 
formula 5, 15, 6, 3, 4+1 aesthetasc, 2+1 
aesthetasc, and 7+1 aesthetasc. Antenna 
(Fig. 2A) 4-segmented, elongate, with ar- 
mature formula 1, 1, 3+1 claw, and 7. First 
and second segments slender, each about 3 
times as long as wide. Seta on these seg- 
ments relatively small and naked. Third 
segment armed with spinules on inner mar- 
gin. Claw on this segment prominent, one 
of three setae very small. Fourth segment 
originating from proximal part of outer 
margin of third segment, wider than long, 
armed terminally with 4 spiniform setae 
and 3 simple setae, outermost seta plumose. 

Labrum (Fig. 2B) reduced, distinctly 
shorter than wide, covering only part of oral 
appendages. Mandible (Fig. 2C) armed with 
1 terminal fleshy, atrophied element tipped 
with a small claw-like process, and 3 long, 
very thin, foliaceous subterminal elements 
covered with minute spinules on all surfac- 
es. Paragnath (Fig. 2D) with 2 hairy lobes 
on medial side. Maxillule (Fig. 2E) taper- 


662 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Foliomolgus cucullus, new genus, new species, female. A, habitus, dorsal; B, anterior part of urosome, 
dorsal; C, urosome, ventral; D, caudal rami, ventral; E, rostral area, ventral; K antennule. Scale bars: A, 0.5 
mm; B, C, 0.2 mm; D—EF 0.1 mm. 


VOLUME 114, NUMBER 3 


ing, proximally with several setules, and 
terminally bilobed, with 5 setae (3 lateral 
ones larger) on anterior lobe and 3 setae on 
posterior lobe. Maxilla (Fig. 2F) 2-seg- 
mented. Basal segment greatly expanded 
proximally, roughly triangular, with 1 ter- 
minal seta. Distal segment terminally trun- 
cated and sclerotized, and armed with 3 un- 
equal lateral setae; terminal portion of distal 
segment covered with numerous, minute 
spinules (Fig. 2G). Maxilliped absent. 

eo (Gis. 2b)wiles 2)(Fie, 3A), lee 3, 
and leg 4 (Fig. 3B) with 3-segmented rami. 
All these legs spiniferous. All legs with 1 
large inner coxal seta; those of legs 1-3 plu- 
mose on one side and weakly spinulated on 
the other side. Inner spine on basis of leg 1 
shorter than first endopodal segment of same 
leg. Posteromedian part of basis of legs 1—4 
ornamented with spinules. Each leg with en- 
dopod distinctly longer than exopod. Formula 
of armature of these legs as follows: 


coxa 0-1: basis 1-—I; 
exp. I-O; I-1; I, 7 
enp. 0-1; O-1; I, 5 


ecm: 


Leg 2 & 3: coxa 0-1: basis 1-0; 


exp. I-0; I-1; II, 7 
enp. O—1; O-—2; III, 3 


Leg 4: coxa Q—1: basis 1—0; 


exp. I-O; I-1; I, 7 
enp. O—1; O—2; III, 2 


Leg 5 2-segmented (Fig. 1B, C). Basal 
segment articulating with fifth pedigerous 
somite, distally armed with 1 small seta and 
spinules. Distal segment elongate, tapering, 
armed with | outer lateral and 3 terminal 
setae; all setae naked, and as long as or 
shorter than distal segment. Leg 6 not seen. 

Male.—Body (Fig. 3C) similar to that of 
female. Length 2.20 mm. Urosome (Figure 
3D) 6-segmented. Genital somite 175 < 317 
zm, distinctly narrower than fifth pedigerous 
somite, without dorsal hood. Genital flap with 
numerous spinules near inner margin. Four 


663 


abdominal somites 154 < 187, 175 x 155, 
150 X 135, and 104 X 113 pm, respectively. 
Caudal ramus 198 X 33 wm (6.0:1), more 
slender than that of female. 

Antennule with 1 additional seta on distal 
portion of third segment. Antenna, labrum, 
paragnath, and maxillule similar to those of 
female. Mandible (Fig. 3E) with more 
prominent claw on tip of terminal element. 
Maxilla (Fig. 3F) with first segment not ex- 
panded, but distal segment as in female. 
Maxilliped (Fig. 3G) 4-segmented. First 
segment with 1 long distal seta. Second 
segment triangular, greatly expanded prox- 
imally, strongly tapering, with a flat, thin 
protrusion at inner proximal corner, and 
armed with 2 rows of thin, truncate spinules 
(one row shorter), 2 rows of epicuticular 
extensions and 2 small setae (one in the 
middle, the other smaller one in distal part) 
on inner margin. Third segment short and 
unarmed. Terminal segment forming a claw 
bearing proximally 2 setae and 1 distal 
membranous process. 

Legs 1—4 as in female. Basal segment of 
leg 5 completely fused with fifth pedigerous 
somite, leaving only 1 distal seta. Free dis- 
tal segment broader and shorter than that of 
female (Fig. 3D). Leg 6 represented by 1 
stiff seta on terminal corner of genital flap. 

Etymology.—The specific name cucullus 
(=hood in Latin) alludes to the hood-like 
dorsal coverture on the fifth pedigerous so- 
mite of the female. 


Discussion 


The genus Hemicyclops best illustrates 
general features of the family Clausidiidae 
(Humes, 1987). The new genus Foliomol- 
gus belongs to the Clausidiidae, because it 
shares with Hemicyclops the important 
characters such as the 7-segmented anten- 
nule, 3-segmented rami on legs 1—4, inner 
spine on the basis of leg 1, 4-segmented 
antenna armed with four and seven ele- 
ments respectively on the third and terminal 
segments, and 5+3 setae on the maxillule. 

Humes & Huys (1992) recognized seven 


664 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


yy, 


Yb, , 
SM ~*~ Wy, 


Fig. 2. Foliomolgus cucullus, new genus, new species, female. A, antenna; B, labrum; C, mandible; D, 
mouth area, including paragnaths; E, maxillule; K maxilla; G, distal part of maxilla; H, leg 1. Scale bars: A—C, 
E-E 0.05 mm; D, 0.02 mm; H, 0.1 mm. 


VOLUME 114, NUMBER 3 665 


=e 


NAY NW 


MAL, 


Roe Zi 
LE Life 
Le 


SSS 
eS 


SSE 


LO EE EE AIS ea 


ANY) Y Ns s % 
ANY JAS] 7 BAAN 
NINN Gc Ys NA BARAN 
ANY Us VARS 
Hy} Nad 9) Mf wy ‘of Ry Ny 
Ni i, VAR ABEN 
V/NINIIN | NANI 4 
Vy —YNSI7 WIN 4 
Ni A\ ARAN 
VIAN WY 


SS 
SSS 


Fig. 3. Foliomolgus cucullus, new genus, new species. Female: A, leg 2; B, leg 4. Male: C, habitus, dorsal; 
D, urosome, ventral; E, mandible; EK maxilla; G, maxilliped. Scale bars: A, B, 0.1 mm; C, 0.5 mm; D, 0.2 mm; 
E, 0.02 mm; FE G, 0.05 mm. 


666 


genera as valid in the Clausidiidae: Clausi- 
dium Embleton, Conchyliurus Bocquet & 
Stock, Doviella Rocha, Hemicyclops Boeck, 
Hippomolgus Sars, Hyphalion Humes, and 
Leptinogaster Pelseneer. In the same year, 
Ho & Wardle (1992) added a new genus 
Pholadicola to this family. Because one of 
these genera, Doviella, is a junior synonym 
of Clausia of the Clausiidae (Kim, 2001), 
the Clausidiidae currently consists of seven 
genera. In the absence of a maxilliped in the 
female and having a claw on the third seg- 
ment of the antenna, Foliomolgus is com- 
parable to the genera Leptinogaster and 
Pholadicola. However the latter two genera, 
both associates of bivalves, carry a 6-seg- 
mented antennule, and fewer setae on the 
antenna, maxillule and legs. 

The morphology of the mandible, bearing 
the rudimentary dorsal (posterior) element 
(claw) and three well-developed foliaceous 
ventral (anterior) elements, is a unique fea- 
ture of the new genus Foliomolgus. As far 
as the Clausidiidae and related families, the 
nereicoliform copepods of Gooding (1963), 
are concerned, the size reduction of mandib- 
ular elements usually involves the ventral 
ones. The extreme case of this armature re- 
duction, where only the dorsal claw-like el- 
ement is retained, is exhibited by some gen- 
era of Catiniidae, Clausiidae, and Synapti- 
philidae. In contrast to this general trend of 
armature reduction, a mandible similar to 
that of the new genus may be seen in co- 
pepodid I of Conchyliurus (see Kim, 1994; 
only three ventral elements are retained in 
the adult) and the adult of Myzomolgus 
where the dorsal claw is reduced and the 
ventral elements are developed. 

In addition to the mandible, the antenna 
also is characteristic within the Clausidiidae 
and related families. In this appendage the 
third segment is produced distally along the 
main axis of the appendage; the fourth seg- 
ment is displaced laterally. 

The transformation of the maxilla in 
which the distal segment is a spatulate lobe, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


one of the diagnostic features of the new 
genus, is a general feature observed in the 
Clausiidae, Nereicolidae and Serpulidicoli- 
dae, the copepods generally associated with 
the polychaetes. Therefore the similarity of 
the maxilla in these copepods may be a 
convergence resulting from the association 
with similar hosts. 


Acknowledgement 


The field survey of this work was sup- 
ported by the Korea Science and Engineer- 
ing Foundation (2000-1-20200-003-3). 


Literature Cited 


Gooding, R. U. 1963. External morphology and clas- 
sification of marine poecilostome copepods be- 
longing to the families Clausidiidae, Clausiidae, 
Nereicolidae, Eunicicolidae, Synaptiphilidae, 
Catiniidae, Anomopsyllidae, and Echiurophili- 
dae. Unpublished Ph.D. thesis, University of 
Washington, Seattle, 276 pp. 

Ho, J.-S. & W. J. Wardle. 1992. Pholadicola intestin- 
alis, new genus and species, a clausidiid cope- 
pod parasitic in a deep-burrowing clam from 
Texas.—Bulletin of Marine Science 51:37—44. 

Humes, A. G. 1987. Copepoda from deep-sea hydro- 
thermal vents.—Bulletin of Marine Science 41: 
645-788. 

, & R. U. Gooding. 1964. A method for study- 

ing the external anatomy of copepods.—Crus- 

taceana 6:238—240. 

, & R. Huys. 1992. Copepoda (Poecilostoma- 
toida and Siphonostomatoida) from deep-sea 
hydrothermal vent areas off British Columbia, 
including Amphicrossus altalis, a new species 
of Erebonasteridae, with note on the taxonomic 
position of the genus Tychidion Humes.—Ca- 
nadian Journal of Zoology 70:1369—-1380. 

Imajima, M., & O. Hartman. 1964. The polychaetous an- 
nelids of Japan, Part If. Allan Hancock Foundation 
Publications, Occasional Paper 26, 452 pp. 

Kim, I.-H. 1994. Copepodid stages of Conchyliurus 
quintus Tanaka, 1961 (Poecilostomatoida, Clau- 
sidiidae) associated with bivalve mollusks.— 
Hydrobiologia 292/293:161—169. 

. 2000. Poecilostomatoid copepods from an in- 

tertidal mud flat in the Yellow Sea.—Journal of 

Natural History 34:367—432. 

. 2001. A new species of Clausia (Copepoda, 

Poecilostomatoida, Clausiidae) associated with 

the polychaete Arenicola brasilliensis Nonata in 

Korea.—Hydrobiologia (in press). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):667—679. 2001. 


Two new species of the Canthocamptus mirabilis group (Copepoda: 
Harpacticoida: Canthocamptidae) from South Korea 


Cheon Young Chang and Teruo Ishida 


(CYC) Department of Biology, Taegu University, Kyungsan 712-714, Korea, e-mail: 
cychang @taegu.ac.kr; 
(TI) 372 Irifunecho, Yoichimachi, Hokkaido 046-0011, Japan 


Abstract.—Two new harpacticoid species belonging to Canthocamptus mir- 
abilis species group, C. odaeensis and C. incurvisetosus, are described from 
South Korea, with an evaluation of the variability of several characters. Both 
species are assumed to be closely related in sharing the apomorphic characters 
of the triangular hyaline membrane on the anal operculum and the male-type 
caudal ramus in females. The occurrence of the C. mirabilis group in South 
Korea is also briefly commented. Canthocamptus incurvisetosus is most widely 
distributed and frequently occurs in mountain waters of South Korea, while C. 
odaeensis is a geographically isolated species. They co-occurred at only one 
location, where the reproductive isolation of these species is reinforced by 
character displacements, especially of the female caudal ramus and the orna- 


mentation of the leg 5 exopod. 


The Canthocamptus mirabilis group, first 
identified as “‘Attheyella-like Canthocamp- 
tus’ by Ito & Takashio (1980), was rec- 
ognized in a series of faunistic studies on 
mountain harpacticoids in Japan (Ishida 
1987, 1989, 1990, 1991; Kikuchi & Ishida 
1988, 1994). It comprises several closely 
related species, occurring with great abun- 
dance and frequency in the various moun- 
tain water bodies of the Far East. Six spe- 
cies are currently known in this group: C. 
mirabilis Stérba, 1968 from China; C. mor- 
imotoi Miura, 1969 from South Korea; and 
C. prominulus Kikuchi, 1993, C. semicir- 
culus Kikuchi, 1993, C. resupinatus Ishida, 
1993, and C. tomikoae Ishida, 1993 from 
Japan. 

In Korea, except for the record of C. 
morimotoi (Miura 1969, Chang 1998), the 
mirabilis group has not been studied in 
spite of its great abundance and frequent 
occurrence in the various mountain water 
bodies. We have fully re-examined the 
specimens belonging to the C. mirabilis 
group which were gathered from 120 lo- 


calities in South Korea since 1986 by the 
senior author. We confirmed that they com- 
prise five species: C. semicirculus Kikuchi; 
an unrecorded species, to be treated later in 
another paper, closely resembling C. mira- 
bilis Stérba from China; C. morimotoi Miu- 
ra; and the two new species described here. 

Collections were made with a dipnet with 
no. 10 mesh. All the specimens were dis- 
sected, drawn, and measured in lactophenol 
on a Cobb’s hole slide. Figures were drawn 
with the aid of a camera lucida. Type ma- 
terial has been deposited in the U.S. Na- 
tional Museum of Natural History, Smith- 
sonian Institution (USNM) and the Natural 
History Museum of Ewha Womans Univer- 
sity, Seoul (EWNMH). 

The caudal ramus, or furca, of species of 
the C. mirabilis group commonly show 
clear sexual dimorphism, but females in 
some populations have a furca similar to the 
males, a reversal of this secondary sexual 
character state (Ito & Takashio 1980, Ishida 
1991). The normal furca and the male-type 
furca of females are abbreviated in the text 


668 


and figure legends to Fn and Fr, respective- 
ly. Abbreviations of enp 1—3 or exp 1-3 are 
used in the description to denote the first to 
third endopodal or exopodal segment of 
ecachwle or 


Canthocamptus odaeensis, new species 
Figs. 1—4 


Material examined.—Twelve 22,7 36, 
foothill spring at Kangnung City (37°45 
SIN, IASI) 23 Seo NOLS. Wee, (Ce 
Y. Chang, including: Holotype 2 (USNM 
310129), allotype 6 (USNM 310130), and 
paratypes (3 22, USNM 310131; 8 22 & 
6 3646, EWNMH 60262). 

Additional material.—Seven 2° °,3 36, 
streamlet at Mt. Odae, 2 Oct 1992, C. Y. 
Chang; 1 2, 3 66, spring at Kujeol-ri, 
Cheongseon, 23 Sep 1993, C. Y. Chang; 5 
22, 3 66, Chodang Cave, Samcheok, 12 
May 1995, H. S. Rho; 5 92, 2 66, same 
as type locality, 21 Jul 1995, C. Y. Chang; 
3 22,1 6, streamlet near Woljungsa Tem- 
ple, Mt. Odae, 28 Jul 1999, J. M. Lee & Y. 
H. Song; 5 22, 6 dd, Mureung Valley, 
Mt. Duta, 9 Oct 1999, J. M. Lee & Y. H. 
Song; 3 2¢, 2 36, trickle near Hwansun 
Cave, Samcheok, 9 Oct 1999, J. M. Lee & 
Y. H. Song; 2 22, 2 6d, Gongsujeon Val- 
ley, Yangyang, 6 Oct 2000, C. Y. Chang. 
All additional material is retained in the re- 
search collection of C. Y. Chang, except 5 
2 @ and 3 36 from Mt. Odae (2 Oct 1992) 
in the collection of T. Ishida. 

Female.—Body (Fig. 1A) length range 
0.64—0.69 mm (0.66 + 0.04 mm, N = 8), 
excluding rostrum and furcal setae; broad- 
est at posterior margin of cephalothorax, ta- 
pering posteriorly. Prosome elliptical. 
Cephalothorax somewhat protruding ante- 
riorly, slightly longer than succeeding 3 
thoracic somites combined. Rostrum not 
prominent, directed anteroventrally, not dis- 
tinct at base. Dorsal and lateral surfaces of 
thoracic somites with few sensilla; posterior 
part of cephalothorax bearing narrow hya- 
line membrane with 10-12 longitudinal 
folds along margin, each bearing 1 sensil- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


lum at its tip. Genital double somite mod- 
erately expanded laterally and slightly 
broadened posteriorly, 1.25 times wider 
than long, subdivided by pair of lateral su- 
tures, each suture with pair of sensilla near 
its dorso-medial end. Genital area T-shaped, 
flanks rather short. Each abdominal somite 
furnished with 1 row of spinules along la- 
terodistal corner, and hyaline fringes with 
crenate posterior margins (Fig. 1B). Disto- 
medial corner of ventral side of anal somite 
bearing 3—4 sharp spinules (Fig. 1C), rela- 
tively undeveloped in comparison with 
those of C. morimotoi (cf. Chang 1998) or 
C. mirabilis sensu Ito & Takashio. Anal 
operculum convex; hyaline membrane (Fig. 
1C) forming obtuse-angled triangle, its tip 
slightly passing dorsal seta of caudal ramus 
in lateral view (Fig. 1B). 

Fr (Fig. 1C) shaped as an inverted bottle; 
about 1.5 times as long as wide, armed with 
1 crescent row of slender spinules along 
medial surface; outer caudal seta ornament- 
ed with 2—3 pairs of sharp secondary spi- 
nules, directed outward and situated from 
proximal 1/5 to 1/3 of the seta (Figs. 1B, 
C); terminal seta stout, slightly bent and 
constricted near base; inner caudal seta 
bare. Fn somewhat suboval (Fig. 1D), about 
1.3 times as long as wide. About 6 spinules 
arranged obliquely at laterodistal corner of 
dorsal surface. Outer terminal seta rather 
straight, not bent, with 3 secondary spinules 
on inner margin and 4—5 acute spinules on 
outer margin. 

Antennule (Fig. 2A) of 8 segments, bear- 
ing | aesthetasc on anterodistal margin of 
fourth article, its tip not reaching distal end 
of antennule. Exopod of antenna (Fig. 2B) 
distinctly 2-segmented; first segment mark- 
edly slender, bearing 1 plumose seta on in- 
ner distal edge with | spinule on laterodistal 
margin; second segment bearing 2+1 plu- 
mose setae. Labrum_ subtriangular, with 
blunt hirsute tip, furnished with 4 subdistal 
spinules. Mandible, maxillule, maxilla, and 
maxilliped (Fig. 2C—F) with typical char- 
acteristics of genus Canthocamptus and 
showing no significant discrepancy from re- 


VOLUME 114, NUMBER 3 


Fig. 1. 


669 


Canthocamptus odaeensis, Female. A, Habitus (dorsal); B, Urosome (lateral); C, Anal somite and 


caudal ramus, showing both ventral (left) and dorsal (right) sides (Fr type); D, Anal somite and caudal ramus 


(Fn type); E, Female leg 5. Scale bars = 0.05 mm. 


lated members of mirabilis group (cf. Ito & 
Takashio 1980, Chang 1998). 

Exopod and endopod of legs 1—4 each 
consisting of 3 segments, except endopod 
of leg 4 with 2 segments. Leg 1 (Fig. 2G) 
enp 1 as long as exopod, with | plumose 
seta on distal 1/5 of inner margin; enp 2 


with | plumose seta on inner distal corner 
and 3 sharp spinules on outer edge; enp 3 
slender, bearing 2 geniculate spines and | 
plumose seta. Exp 1 longer than next two 
segments; exp 2 with | inner seta and 1 
outer spine; exp 3 bearing 2 outer spines 
and 2 geniculate setae. Distal edge of inter- 


670 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


| ney 


sf 


/yy\ 


Fig. 2. Canthocamptus odaeensis, Female. A, Antennule; B, Exopod of antenna; C, Mandible; D, Maxillule; 
E, Maxilla; EK Maxilliped; G, Leg 1; H, Leg 2. Scale bars = 0.05 mm. 


VOLUME 114, NUMBER 3 


Fig. 3. 


coxal sclerites of legs 2—4 with several spi- 
nules. Ornamentation of legs 2—4 as follows 
(in armature formula, Roman numerals in- 
dicate spines and Arabic numerals represent 
setae): 


Leg 2: basis I-O; exp I-0; I-1; Il, 2, 1 
enp 0-1; 0-1; I, 2, 2 
Leg 3: basis 1—0; exp I-0; I-1; III, 2, 2 
enp 0-1; 0-1; I, 2, 2 
Leg 4: basis 1—0; exp I-O; I-1; III, 2, 2 


enp O-1; I, 2, 2 
Distal end of leg 5 baseoendopod (Fig. 


671 


Canthocamptus odaeensis. A, Female leg 3; B, Female leg 4; C, Male leg 4. Scale bars = 0.05 mm. 


1E) not reaching middle of exopod; bas- 
eoendopod not confluent at its base, con- 
nected by intercoxal sclerite; bearing 6 spi- 
niform setae, of which outermost two setae 
shortest and next innermost seta longest. 
Exopod rather oblong, slightly tapering dis- 
tally, about 2.7 times as long as broad; inner 
margin slightly swollen; armed with 5 se- 
tae, distalmost seta of which slightly longer 
than exopod; 2 spinules usually present on 
distomedial corner on ventral side of exo- 
pod near base of inner seta. 

Male.—Body (Fig. 4A) length from 
0.63—0.68 mm (0.64 + 0.04 mm, N = 6). 
Overall appearance like female, except with 


672 


more slender caudal ramus. Hyaline mem- 
brane on anal operculum (Fig. 4B) trian- 
gular with convex margin, its tip reaching 
level of dorsal seta. Leg 2 (Fig. 4C), enp 1 
with | inner seta; enp 2 bearing 3 inner, 2 
long terminal setae, with outer spinules; 2 
inner setae of enp 2 and inner seta of exp 
3 modified with pectinate tip. Leg 3 (Fig. 
4D), enp 2 armed with | acute spiniform 
seta and | process (apophysis) with termi- 
nal barbs, not extending beyond exp 3; enp 
3 elongate, slightly longer than combined 
lengths of two proximal segments, tapering 
distally, ending with 2 plumose setae; exp 
2 armed with stout spiniform process on 
outer distal corner, its tip usually not reach- 
ing or rarely reaching slightly beyond distal 
margin of exp 3; exp 3 with 2 inner setae, 
distal one bent with pectinate tip. Leg 4 
(Fig. 3C) nearly same shape as that of fe- 
male, including outer terminal seta on exp 
3 armed with more than 18 spinules on out- 
er margin of seta; distal inner seta of exp 3 
modified with pectinate tip. Baseoendopo- 
dal lobe of leg 5 (Fig. 4E) triangular, not 
reaching middle of exopod; bearing 2 spi- 
niform setae, inner seta about 1.3 times lon- 
ger than outer; inner margin bare. Exopod 
2.1 times longer than broad, armed with 6 
setae in total, consisting of 1 weak plumose 
seta at about middle of medial margin, | 
inner distal, 2 terminal, and 2 outer spini- 
form setae. Leg 6 (Fig. 4B) represented by 
small plate bearing 1 spine and 2 slender 
setae. 

Variability.—Females with normal-type 
caudal ramus (Fn) were rarely observed 
(fewer than 1% of all specimens examined). 
No significant variation was observed in the 
caudal ramus among the specimens exam- 
ined, except that about 8% of Fr lacked the 
medial spinules (while all males examined 
had the spinules). The spiniform process on 
the male leg 3 exp 2 usually did not reach 
or reached only slightly beyond the tip of 
the leg 3 exp 3. Most females (78% of spec- 
imens examined) had two spinules on the 
inner distal edge of the ventral surface of 
the leg 5 exopod. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


The ornamentation of the outer terminal 
caudal seta was characteristic and consis- 
tent in all the specimens examined, as was 
the unmodified outer terminal seta (armed 
with about 18 or more secondary spinules 
on outer margin of the seta) on the male leg 
4 exp 3. No particular difference was de- 
tected in the major ornamentation of legs 
2-4. 

Etymology.—The specific name is taken 
from Mt. Odae, because the distribution of 
this new species is confined to Mt. Odae 
and the surrounding area. 

Remarks.—Affinities with related species 
are discussed together with the next species. 


Canthocamptus incurvisetosus, new 
species 
Figs. 5—6 


Material examined.—Eleven 22,5 36, 
mountain streamlet in Seonamsa Valley, 
Mt. Jogye (34°59'26"N, 127°19'53"E), 13 
Jan 2000, leg. C. Y. Chang, J. M. Lee & Y. 
H. Song, including: holotype 2 (USNM 
310132), allotype 6 (USNM 310133), and 
paratypes (3 22, USNM 310134;7 22 & 
4 363, EWNMH 60261). 

Additional material.—Eleven 22, 9 
66, spring, Ssangyesa Temple, Mt. Chiri, 
21 Jan 1987, C. Y. Chang; 3 2 2, streamlet, 
Mt. Wolchul, 7 Feb 1987, C. Y. Chang; 9 
22,4 3d, spring, Mt. Namdogyu, 8 Aug 
1987, C. Y¥. Chang: 5 99° 4°66 micklemn 
Seonamsa Valley, Mt. Jogye, 31 Jan 1991, 
CY. Chang; 3° 2 2) 2°o 5, Posereseane 
Ulung I, 3 Mar 1995, C. Y. Chang; 3 dd, 
streamlet, Mt. Chiak, 27 Jun 1995, C. Y. 
Chang; 5 22, 4 d6, streamlet, Mt. Sorak, 
3 May 1998, S. M. Yoon; 4 22, 3 dd, 
trickle in Naewonsa Valley, Mt. Chiri, 26 
Oct 1999, C. Y. Chang & J. M. Lee; 10 2 &, 
6 36, Mt. Seongju, Poryong, 20 Nov 1999, 
C. Y. Chang. All additional material is re- 
tained in the research collection of C. Y. 
Chang, except 3 22 and 2 66d from Mt. 
Chiri (21 Jan 1987) in the collection of T. 
Ishida. 

Female.—Body length 0.69—0.73 mm 


VOLUME 114, NUMBER 3 673 


—— 


Fig. 4. Canthocamptus odaeensis, Male. A, Habitus (dorsal); B, Urosome showing both ventral (left) and 
dorsal (right) sides; C, Leg 2; D, Leg 3; E, Leg 5. Scale bars = 0.05 mm. 


674 


(0.71 + 0.02 mm, N = 7), excluding ros- 
trum and caudal setae; overall appearance 
and mouthparts not showing significant dif- 
ferences from those of preceding species. 
Each abdominal somite bearing | row of 
spinules along laterodistal margin, and 
adorned with crenate hyaline frill (Fig. 5A). 
Posteromedial corner of ventral side of anal 
somite bearing 4—5 acute spinules. Anal 
operculum convex, furnished with hyaline 
membrane (Figs. 5B, C) forming obtuse-an- 
gled triangle with convex margin, but its tip 
projecting sharply. 

Caudal ramus of normal type (Fig. 5B) 
ovoidal, about 1.76 times as long as wide, 
ornamented with crescent row of slender 
spinules along medial surface; laterodistal 
part of dorsal surface adorned with 6—9 spi- 
nules; dorsal keel over dorsal seta not 
prominent; outer terminal caudal seta twist- 
ed and curved inward, ornamented with 2— 
3 long, inward-directed secondary spinules 
on medial face of the seta; terminal seta 
slightly bent and constricted near base; in- 
ner caudal seta bare and strongly bent near 
its base. Fr (Fig. 5C) shaped as inverted 
bottle, about 1.5 times as long as wide, 
without spinules on outer distal part of dor- 
sal surface; with or without crescent row of 
slender spinules along medial surface, this 
row sometimes present on only one ramus; 
other characters same as those of normal 
type ramus. 

Segmentation and ornamentation of legs 
1—4 exactly as those of preceding species. 
Leg 5 (Fig. 5D) exopod elongate, a little 
tapering distally, about 3.2 times as long as 
broad; inner margin slightly swollen; armed 
with 5 setae, of which inner distal seta al- 
ways slightly shorter or as long as exopod; 
inner seta located near distomedial edge of 
exopod; spinules on distomedial corner of 
exopod usually absent, but sometimes 2—3 
spinules present. Baseoendopod reaching 
about %4 length of exopod; baseoendopod 
not confluent at its base, connected by in- 
tercoxal sclerite; bearing 6 spiniform setae, 
two outermost setae smallest and two in- 
nermost longest. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Male.—Body (Fig. 6A) length from 
0.54—0.63 mm (0.59 + 0.05 mm, N = 5). 
Hyaline membrane of anal operculum (Fig. 
6B) triangular, its tip extending beyond lev- 
el of dorsal setae of the caudal ramus. Dis- 
tomedial corner of anal somite bearing 3—4 
spinules, these relatively stronger than cor- 
responding spinules of female. Caudal ra- 
mus (Fig. 6B) rather elongate, tapering pos- 
teriorly, about 1.76 times as long as wide, 
with no spinule row along medial surface; 
dorsal keel not prominent and dorsal sur- 
face smooth except for dorsal seta; outer 
terminal caudal seta normal, i.e., not twist- 
ed and not curved inward as in female, and 
ornamented with 2—3 outwardly directed 
secondary spinules; inner caudal seta un- 
armed. Leg 3 (Fig. 6D) exp 2 armed with 
stout spiniform process on outer distal cor- 
ner, its tip extending far beyond distal end 
of exp 3. Leg 4 (Fig. 6E) exp 3 outer ter- 
minal seta with 3—4 sharp and strong spi- 
nules on outer margin. Other characters of 
legs 2—4 not showing significant discrep- 
ancies from preceding species. Baseoendo- 
pod of leg 5 (Fig. 5E) protruding, but not 
reaching 4% of exopod; bearing 2 terminal 
spiniform setae, inner seta about 1.43 times 
longer than outer seta; inner margin bare. 
Exopod 2.4 times longer than broad, armed 
with 6 setae in total, these consisting of 1 
weak plumose seta on inner middle, 1 inner 
distal, 2 terminal, and 2 outer spiniform se- 
tae. Leg 6 (Fig. 6F) a small plate bearing 1 
stout spine and 2 slender setae; spine 1.4 
times longer than next seta. 

Variability—Fn always possesses the 
spinule array on the medial surface of the 
caudal ramus, while 38% of Fr (male-type 
caudal ramus) were unarmed, as were more 
than 95% of the males. 18% of Fr had spi- 
nules on one caudal ramus but not on the 
other. The spinule arrangement on the outer 
distal corner was absent in Fr, and some- 
times also absent in Fn; rarely individuals 
possessed both the different types of orna- 
mentation. Usually (in more than 95% of 
cases) the spiniform process on the outer 
distal edge of the male leg 3 exp 2 much 


VOLUME 114, NUMBER 3 675 


YW, 
Gyn 


van 1) 


fos] 


Fig. 5. Canthocamptus incurvisetosus. A, Urosome of female (Fn type, ventral); B, Anal somite and caudal 
ramus of female (Fn type, dorsal); C, Anal somite and caudal ramus of female (Fr type, dorsal); D, Female leg 
5; E, Male leg 5. Scale bars = 0.05 mm. 


676 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Canthocamptus incurvisetosus, male. A, Habitus (dorsal); B, Anal somite and caudal ramus, showing 
both ventral (left) and dorsal (right) sides; C—E, legs 2—4; FE leg 6. Scale bars = 0.05 mm. 


VOLUME 114, NUMBER 3 


exceeded the tip of leg 3 exp 3. Only 28% 
of specimens examined had 2-3 spinules on 
the inner distal edge of the ventral surface 
of the female leg 5 exopod. 

All individuals examined had the typical 
shape of twisted and inward-curved outer 
terminal caudal setae as described and il- 
lustrated above, with 2—3 secondary spi- 
nules, or exceptionally with 4 spinules. The 
outer terminal seta on male leg 4 exp 3 was 
also consistently furnished with 3—4 sharp 
strong spinules on its outer margin. No par- 
ticular difference was detected in the major 
ornamentation of legs 2—4. 

Etymology.—The proposed specific 
name is taken from the Latin incurvus (bent 
inward) and setosus (bearing seta), which 
alludes to the possession of the twisted and 
inward-curved outer terminal seta of the fe- 
male caudal ramus, the distinctive charac- 
teristic of this species. 

Remarks.—In having the apomorphic 
character-combination of the triangular hy- 
aline membrane on the anal operculum and 
the male-type caudal ramus of the female, 
both these new species differ from C. sem- 
icirculus Kikuchi, C. tomikoae Ishida, C. 
mirabilis Stérba, and an undescribed spe- 
cies from Korea. They share these derived 
states with C. morimotoi Miura, C. resupi- 
natus Ishida and C. prominulus Kikuchi. 
The affinities among the former group of 
four species will be discussed elsewhere, 
when C. mirabilis Stérba is redescribed on 
the basis of the newly collected material 
from type locality near Beijing, China, to- 
gether with the description of another new 
Korean species. 

Canthocamptus odaeensis n. sp. shares 
with C. morimotoi a well-developed trian- 
gular hyaline membrane on the anal oper- 
culum and the normal ornamentation of the 
outer distal seta of male leg 4 exopod 3 
G.e., not modified with 3—4 sharp and 
strong secondary spinules on outer margin, 
but armed with about 18 or more spinules 
on outer margin). However, females of C. 
odaeensis may express one of two different 
kinds of caudal ramus, the normal-type fe- 


677 


male caudal ramus has a spinule row near 
the distolateral corner of dorsal surface, and 
the male caudal ramus has medial spinules. 
Females of C. morimotoi express only one 
kind of caudal ramus, which lacks the spi- 
nule row near the distolateral corner of dor- 
sal surface, and the male caudal ramus has 
smooth medial face without medial spi- 
nules. However, it is distinct in showing Fr 
while the caudal ramus of both sexes are 
subconical in C. morimotoi. In possessing 
the terminal seta of leg 5 slightly longer 
than leg 5 exopod and the medial spinules 
on the caudal ramus of the male, C. odaeen- 
sis 18 similar to C. resupinatus, but is dis- 
tinguished from that species by the normal 
outer distal seta on the male leg 4 exopod 
3, and the shape of the outer terminal cau- 
dal seta. 

Canthocamptus odaeensis resembles C. 
prominulus Kikuchi in having a relatively 
short spiniform process on the outer distal 
edge of the male leg 3 exopod 2, but the 
former species differs from the latter in 
having an unmodified terminal seta on the 
male leg 4 exopod 3, and in possessing the 
terminal seta on female leg 5 exopod slight- 
ly longer than the exopod. Canthocamptus 
odaeensis is also distinguished from C. in- 
curvisetosus by the shape of the outer ter- 
minal caudal seta, the presence of a setule 
array on the medial surface of male caudal 
ramus, and the relative length of the female 
leg 5 exopod with its terminal seta. 

Canthocamptus incurvisetosus closely re- 
sembles C. resupinatus in the hyaline mem- 
brane of the anal operculum and in the or- 
namentation of the outer distal seta of the 
last exopodal segment of the male leg 4, 
together with the twisted and inward- 
curved outer caudal setae. It differs from C. 
resupinatus in having the male caudal ra- 
mus with a smooth medial surface and the 
female caudal ramus with several spinules 
near distolateral corner of dorsal surface, as 
well as in having the terminal seta on the 
exopod of the female leg 5 shorter than the 
ramus itself. The inward curvature of the 
outer terminal caudal setae of C. incurvi- 


678 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


o 


126°E 


128°E 130°E 


Fig. 7. Occurrence of species of the Canthocamptus mirabilis group in South Korea. Symbols as follows: 
@ Canthocamptus odaeensis, * Canthocamptus semicirculus, 0 Canthocamptus incurvisetosus, * Canthocamptus 


sp., A Canthocamptus morimotoi. 


setosus and C. resupinatus is the decisive 
feature distinguishing these two species 
from other congeners. These setae are the 
site of initial male mechanical contact with 


the female during mating of the C. mirabilis 
group. The male always grasps the proxi- 
mal portion of female’s caudal setae (cf. 
Chang 2001, Fig. 4A, B). The ornamenta- 


VOLUME 114, NUMBER 3 


tion of the setae is somewhat different 
among the members of the species group, 
as indicated in the specific names, the in- 
ward curvature of the outer terminal caudal 
setae of C. incurvisetosus and C. resupi- 
natus. However, morphological details of 
these setae differ between the two species: 
smoothly curved with 2—3 secondary spi- 
nules on the medial surface of the seta of 
C. incurvisetosus, and somewhat sharply 
bent like a sickle with 7 setiform spinules 
of C. resupinatus. 

Four of five species occurring in South 
Korea are endemic, and only C. semicir- 
culus Kikuchi is known to be widely dis- 
tributed in Japan (below 37°N) and Taiwan 
(Kikuchi & Ishida 1994). Canthocamptus 
incurvisetosus n. sp. frequently occurs in 
mountain waters such as streamlets, trick- 
les, and springs in South Korea (Fig. 7). It 
was collected from 63 of the 120 localities 
studied, and ranged throughout South Ko- 
rea except where C. odaeensis was found. 
Thus it co-occurred with C. morimotoi over 
the area from Mt. Sorak (northeastern South 
Korea) to Mt. Joryeong (central South Ko- 
rea), with C. semicirculus along the Sobaek 
Mountains (which run diagonally south- 
westward), and with an unrecorded species 
of the mirabilis group in western South Ko- 
rea. Like C. resupinatus in Japan, C. 
odaeensis is geographically restricted. It is 
found within narrow limits from south of 
Mt. Sorak, the northern limit of its range, 
to north of Mt. Taebaek, the southern limit 
(from 37°15’ to 38°10’ N), along the eastern 
slope of the Taebaek Mountains. This spe- 
cies co-occurred with C. incurvisetosus and 
C. morimotoi at only one location, Han- 
gyeryong of Mt. Sorak at the northern bor- 
der of its range. There, all female C. 
odaeensis were Fn type while all C. in- 
curvisetosus were Fr type, and C. incurvi- 
setosus possessed the elongated female leg 
5 exopod bearing one seta on the medial 
edge as in the male, as well as several fine 
setules as in C. semicirculus. This suggests 
that reproductive isolation among these spe- 
cies where they co-occur is reinforced by 


679 


character displacement, especially of the fe- 
male caudal ramus and the ornamentation 
of the leg 5 exopod. 


Acknowledgments 


We are grateful to Dr. Janet Reid for 
reading the manuscript with the helpful 
comments. We also wish to pay special 
thanks to several anonymous reviewers who 
made valuable suggestions for improve- 
ment of the manuscript. This research was 
supported by a Taegu University Research 
Grant, 2000. 


Literature Cited 


Chang, C. 1998. Redescription of Canthocamptus mor- 
imotoi Miura, a stygobiontic harpacticoid spe- 
cies from Korea, with a brief review on C. mir- 
abilis group.—Korean Journal of Biological 
Sciences 2:427—434. 

. 2001. Redescription of Canthocamptus mira- 
bilis Stérba, based on the topotypic material 
from China.—The Korean Journal of System- 
atic Zoology 17:1—11. 

Ishida, T. 1987. Freshwater harpacticoid copepods of 
Hokkaido, northern Japan.—Scientific Reports 
of the Hokkaido Salmon Hatchery 41:77-119. 

. 1989. Copepods in the mountain waters of 

Honshu, Japan.—Scientific Reports of the Hok- 

kaido Salmon Hatchery 43:1—21. 

. 1990. Copepods in the mountain waters of 

Kyushu, Tsushima and Ryukyu Islands, south- 

western Japan.—Scientific Reports of the Hok- 

kaido Salmon Hatchery 44:39—51. 

. 1991. Variation in the species of freshwater 
harpacticoid copepods in Japan. I. Canthocamp- 
tus mirabilis Stérba—Bulletin of Plankton So- 
ciety of Japan, Special Volume 1991:391—396. 

Ito, T., & T. Takashio. 1980. Canthocamptus mirabilis 
Stérba (Copepoda, Harpacticoida) from Hok- 
kaido, northern Japan.—Annotationes Zoologi- 
cae Japonenses 53:210-—219. 

Kikuchi, Y., & T. Ishida. 1988. On some freshwater 
harpacticoids from Japan, closely related to 
Canthocamptus mirabilis Stérba.—Hy drobiolo- 
gia 168/169:401—407. 

, & . 1994. A species group of genus 
Canthocamptus (Copepoda: Harpacticoida) in 
Japan, including descriptions of four new spe- 
cies.—Bulletin of the Biogeographical Society 
of Japan 49:37—46. 

Miura, Y. 1969. Results of the speleological survey in 
South Korea 1966 XIV. Subterranean harpacti- 
coid copepods of South Korea.—Bulletin of the 
National Science Museum, Tokyo 12:241—254. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):680—685. 2001. 


Exopod and protopodial endite III of the maxillae of Sarsiellinae 
(Crustacea: Ostracoda: Myodocopa) 


Louis S. Kornicker 


Department of Systematic Biology, National Museum of Natural History, Smithsonian Institution, 
Washington, D.C. 20560-0163, U.S.A. 


Abstract.—The number of bristles on the exopod and, also, the number and 
distribution of bristles and claws on protopodial endite III of the maxillae of 
species of the Sarsiellinae (Ostracoda) are of value in discriminating species, 
and should be described and illustrated in descriptions of new species. 


Poulsen (1965:46) stated, ““Most of the 
species of the family Sarsiellidae differ 
from one another in only a few characters 
and the differences themselves are not very 
conspicuous.’ The purpose of the present 
study is to evaluate the possible importance 
in taxonomic discrimination within the Sar- 
siellinae of two characters of the maxilla: 
1, the number of bristles on the exopod; and 
2, the number and distribution of bristles 
and claws on endite III of the protopod. 

The number of exopodial bristles gener- 
ally have been noted in species descriptions 
enabling most of the data on exopodites in 
Table 1 to be obtained from a survey of the 
literature; however, the number and distri- 
bution of bristles and claws on endite III of 
the protopod seldom have been noted in de- 
scriptions, requiring that most of the data 
on endite III in Table 1 had to be obtained 
from examination of specimens. 

The maxilla of the Sarsiellinae has a pro- 
topod with three endites: endite I, endite II 
(middle), endite II (distal) (Fig. 1B, D). 
Distally, the limb has a 2-jointed endopod, 
and a single jointed exopod (Fig. 1A). The 
maxilla of the adult male is reduced (Fig. 
KE). 

When the maxilla is viewed in its natural 
position on the body, the basis and endopod 
have their broadest dimensions parallel to 
the carapace. The 2nd endopodial joint is 
curved inward towards the mouth. The 
short exopod is on the outer side of the limb 


just posterior to the Ist endopodial segment. 
The endites form a diagonal row medial to 
the endopod with endite III lateral and 
slightly posterior to endites I and II. Be- 
cause of its lateral location, the distribution 
of bristles on endite III is generally visible 
when the limb is viewed from the outside. 


Exopod 


With the exception of some members of 
the Cylindroleberididae, which have max- 
illae with a quite different morphology than 
other families of the Myodocopina, the 
maxillae of the Myodocopina are biramous. 
The exopods of the maxillae of members of 
the families Cypridinidae, Philomedidae, 
and Rutidermatidae bear three bristles; 
whereas, the exopods of the Sarsiellidae 
bear either | (rarely), 2 (Fig. 2G), or 3 bris- 
tles (Fig. 2A—C, F) (Table 1). 

The ontogeny of only a few species of 
Sarsiellinae are known. Three species (Sar- 
siella japonica Hiruta, 1977; Sarsiella mis- 
akiensis Kajiyama (Hiruta, 1978), and Eu- 
sarsiella ryanae Kornicker & Iliffe, 2000) 
have the same number of exopodial bristles 
on Instar I and the adult. Juveniles of both 
sexes have maxillae similar in type to that 
of the adult female. 

The exopod of the adult male is smaller 
than that of the female and a lobe may be 
absent; however, the bristles of the exopod 
are well developed and ringed. Of the 28 


VOLUME 114, NUMBER 3 


681 


Table 1.—Number of species in genera of Sarsiellinae having either 2 or 3 exopodial bristles and 4, 5, or 6 
bristles and claws on endite III of the protopod. (The number of exopodial bristles is mostly from adult females, 
but includes a few adult males. The number of bristles on endite III is mostly from adult females, but includes 
a few instar IV males and females, which are believed to have the same number of bristles as on the adult; the 


list does not include all species in the subfamily.) 


Exopod 
Genus 2 br. 3 br. 


Adelta 1 
Alphasarsiella —: 
Ancohenia —_ 
Anscottiella = 
Chelicopia 6 
Cymbicopia — 
Eurypylus 3 
Eusarsiella 25 
Junctichela 1 
Metasarsiella 0 
Neomuelleriella 2) 
Parasarsiella 0) 
Sarsiella 13 
Spinacopia 5 
Tetrasarsiella — 


WL oOoMnNR REN © 


species considered herein of which adult 
males and females are known, 26 have the 
same number of exopodial bristles (two or 
three). The males of two species have three 
exopodial bristles compared to two on the 
females. 

Adult females and juveniles of the Sar- 
siellinae are voracious predators, whereas, 
adult males are not. This is reflected in the 
reduced maxilla of the adult male (Fig. 1C); 
the reduction of the maxilla suggests that it 
is not used in feeding. The well developed 
exopodial bristles on the adult male maxilla 
suggest that they are mainly used for sens- 
ing. 

If the species of the Cypridinidae, Phi- 
lomedidae, and Rutidermatidae, were to be 
considered as outgroups, three bristles 
could be interpreted as the plesiomorphic 
character state. Then, the two bristles on 
some exopods of species of Sarsiellinae 
could be interpreted to be the apomorphic 
character state. However, because both two 
and three exopodial bristles occur on di- 
verse genera of the Sarsiellinae, I interpret 
the loss of one bristle to be the result of 
convergence or parallelism, and probably 


Endite II 

4 br. 5 br. 6 br. 
_ = 1 
— = 1 
L5 y) a 
_ = 3 
as a 1 
aaa 3 1 

3 4 34 

1 Le 1 
aed 1 =3 
an = 3 
a za 1 
— 4 
— — 14 


not of significance in discriminating genera 
of Sarsiellinae as presently defined. 

In conclusion, the number of bristles on 
the exopod appears to be a useful character 
in defining some species. 


Protopodial Endite II 


The bristles of endite III of the reduced 
maxilla of the adult male are weakly de- 
veloped and not considered further herein. 
Endite III of the adult female and juveniles 
of both sexes is broad and globose with 
well developed bristles. When the endite is 
flattened under a cover slip, the bristles 
generally appear to be along, or close to, an 
edge of the endite (Fig. 2). Endite III of the 
species studied herein bears a total of 4, 5, 
or 6 bristles and claws (Table 1). In the dis- 
cussion below claws and bristles are not 
differentiated, both are included as bristles. 

The number of bristles on endite III of 
juvenile instars was determined for only a 
few species (Table 2). The four species of 
instar I examined all have 4 bristles; the 
remaining instars and the adult females ex- 
amined have a total of 4 to 6 bristles and 


682 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


—— <—___ 


Fig. 1. Eusarsiella ryanae Kornicker and Iliffe: Adult female, holotype, USNM 194475. A, left maxilla, 
lateral view (endites I and II and 3 bristles of endite II] not shown); B, endites left maxilla, medial view; C, 
adult male, paratype, USNM 194476, right maxilla, lateral view; D, stylized lateral view of female left maxilla. 
Illustrations A—C adapted from Kornicker and Iliffe (2000, figs. 18c, e and 20a, respectively.) (bas = basis; cx 
= coxa; end = endopod; ex = exopod; arrows indicate anterior; Roman numerals identify endites.) 


claws. The species Junctichela pax has 4 _ stars are known for only a few species. En- 
bristles on all stages. The remaining species dite III of instar IV and the adult female 
examined have 5 or 6 bristles on instars I— have the same number of bristles, except 
IV and the adult female, but the early in- for endite III of Eusarsiella radiicosta, 


VOLUME 114, NUMBER 3 683 


Fig. 2. Endite Il of maxilla of species of Sarsiellinae: A, Eusarsiella dispar Kornicker, 1986, USNM 158033, 
adult female, left limb, lateral view; B, Eusarsiella maurae (Kornicker, 1977), USNM 156739, adult or A-1 
female, left limb, lateral view; C, Eurypylus hapax Kornicker & Iliffe, 2000, USNM 194494, instar IV male, 
right limb, lateral view; D, Anscottiella vertex Kornicker, 1991, USNM 159317, adult female, left limb, medial 
view; E, Ancohenia robusta (Brady, 1890), USNM 193623, instar III female, left limb, lateral view; EK Spinacopia 
menziesit Kornicker, 1969, USNM 122086, adult female, right limb, lateral view; G, Eusarsiella styx Kornicker 
& Iliffe, 1989, USNM 193367, adult female, left limb, lateral view. (Arrows indicate anterior.) 


684 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Number of bristles (includes claws) on endite III of the protopod of stages of selected species of 
Sarsiellinae (both sexes are combined for juvenile instars). (nd = no data) 


Stages 
Species I Il Ill IV Adult female 
Ancohenia robusta nd nd 5 nd nd 
Chelicopia obex nd nd nd 6 nd 
Cheliocopia radix nd nd nd 6 nd 
Eurypylus hapax nd nd nd 5 nd 
Eusarsiella capillarus nd nd nd 6 nd 
Eusarsiella merx 4 =) nd 5 nd 
Eusarsiella radiicosta nd nd nd 5—6* 6 
Eusarsiella ryanae 4 6 6 6 6 
Eusarsiella thrix nd nd nd 6 nd 
Junctichela pax 4 4 4 4 4 
Sarsiella capsula nd nd nd 6 6 
Spincaopia sandersi 4 nd nd 6 6 


* Specimen with 5 bristles on one limb and 6 on other limb. 


which has 5 bristles on one limb of instar 
IV and 6 on the other; that species has 6 
on the adult female. Because of the simi- 
larity in the number of bristles on endite III 
of both instar IV and the adult female, e1- 
ther stage is included in Table 1. 

The number of bristles on endite III of 5 
adult females and 2 instars IV of Spinaco- 
pia sandersi were counted. All had 6 bris- 
tles. Because of this it is tentatively as- 
sumed in this paper that the number of bris- 
tles is fairly constant in those two stages in 
species in the Sarsiellinae. 

Endite III of seven species in five genera 
of Sarsiellinae are illustrated in Fig. 2. All 
were drawn with the limb flattened under a 
cover slip. Endite III is usually broader than 
either endites I and II. Endite III bears three 
to five ringed bristles and one or two unr- 
inged pectinate claws. On some species the 
anterior bristles of endite III are on a pro- 
jection (Fig. 2D, E, G). The anterior bristles 
extend anteriorly towards the mouth, and 
the anterior of these is generally longer than 
other bristles (Fig. 2A—C). Claws are locat- 
ed between the anterior and posterior 
groups of ringed bristles and each curves 
anteriorly (Fig. 2). On some species two or 
three bristles form anterior and posterior 
groups separated from each other by a wide 
space (Fig. 2A—C, G). 


In conclusion, the distribution of bristles 
and claws on endite III, as well as the 
lengths of bristles, vary considerably 
among species of the Sarsiellinae, and may 
be a valuable character for discriminating 
species. Therefore, I highly recommend that 
endite III be illustrated in species descrip- 
tions of the Sarsiellinae. 


Acknowledgments 


I thank Molly K. Ryan for inking the ap- 
pendages in Fig. 2, and Elizabeth Harrison- 
Nelson (Smithsonian Institution) for assist- 
ing in preparation of the paper. 


Literature Cited 


Brady, G. S. 1890. On Ostracoda collected by H.B. 
Brady, Esq., L.L.D.ER.S., in the South Sea Is- 
lands.—Transactions of the Royal Society of 
Edinburg 35 (part 2, number 14):489—525. 

Hiruta, S. 1977. A new species of the genus Sarsiella 
Norman from Hokkaido, with reference to the 
larval stages (Ostracoda: Myodocopina).—Jour- 
nal of the Faculty of Science, Hokkaido Uni- 
versity 6 (Zoology) 21:44—60. 

. 1978. Redescription of Sarsiella misakiensis 
Kajiyama from Kokkaido, with reference to the 
larval stages (Ostracoda: Myodocopina).—Jour- 
nal of the Faculty of Science, Hokkaido Uni- 
versity 6 (Zoology) 21:262—278. 

Kornicker, L. S. 1969. Morphology, ontogeny, and in- 
traspecific variation of Spinacopia, a new genus 


VOLUME 114, NUMBER 3 


of myodocopid Ostracod (Sarsiellidae).— 
Smithsonian Contributions to Zoology 8:1—50. 
. 1977. Sarsiella maurae, a new species of ma- 
rine Ostracoda (Sarsiellidae: Myodocopina) 
from Bahia de los Angeles, Gulf of California, 
Mexico.—Proceedings of the Biological Socie- 
ty of Washington 90:676—684. 

. 1991. Myodocopid Ostracoda of Enewetak 
and Bikini Atolls.—Smithsonian Contributions 
to Zoology 505:1—140. 

, & T. M. Iliffe. 1989. Ostracoda (Myodocopi- 


685 


na: Cladocopina, Halocypridinia) mainly from 

anchialine Caves in Bermuda.—Smithsonian 

Contributions to Zoology 475:1—-88. 

, & . 2000. Myodocopid Ostracoda from 
Exuma Sound, Bahamas, and from marine 
caves and Blue Holes in the Bahamas, Bermu- 
da, and Mexico.—Smithsonian Contributions to 
Zoology 606:1—98. 

Poulsen, E. M. 1965. Ostracoda-Myodocopa, 1: Cy- 
pridiniformes-Rutidermatidae, Sarsiellidae and 
Asteropidae.—Dana Report 65:1—484. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):686—693. 2001. 


Cypridina olimoblonga Kornicker, a new name for the junior primary 
homonym Cypridina oblonga Jones & Kirkby, 1874, and clarification 
of its authorship, and the authorship of the Palaeozoic genera 
Cypridinella, Cypridellina, Sulcuna, Rhombina, and Offa 
(Crustacea: Ostracoda) 


Louis S. Kornicker 


Department of Systematic Biology-IZ, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A. 


Abstract.—A new name, Cypridina olimoblonga Kornicker, is proposed for 
the junior primary homonym Cypridina oblonga Jones & Kirkby 1874 (non 
C. oblonga Grube 1859), type species of Sylvesterella Kornicker & Sohn 2000. 
Authorship of the replaced species, as well as other species described as new 
in ““A Monograph of the British Fossil Bivalved Entomostraca from the Car- 
boniferous Formations” (1874), should be attributed to T. Rupert Jones & 
James W. Kirkby, not to Jones, Kirkby & Brady. It is concluded that the correct 
authorship of the genera Cypridinella, Cypridellina, Sulcuna, Rhombina, and 
Offa should be attributed to T. Rupert Jones. 


Kempf (in litt., 24 Aug 2000) informed 
me that the species listed in Kornicker & 
Sohn (2000:20) as Cypridina oblonga Jones 
& Kirkby 1874, type species of Sylvester- 
ella Kornicker & Sohn 2000, is a junior pri- 
mary homonym of Cypridina oblonga Gru- 
be 1859. The illustrations of the species by 
Grube (1859: pl. 12: figs. 2-5) clearly iden- 
tify it as a member of the Cylindroleberi- 
dinae. Therefore, I herewith rename Cypri- 
dina oblonga Jones & Kirkby 1874, as Cy- 
pridina olimoblonga Kornicker nom. nov. 
Olim is from the Latin meaning “formerly” 
or “‘once’’. 

In the same letter, Kempf (in litt., 24 Aug 
2000) stated, ““Authors of this species [the 
junior homonym Cypridina oblonga] are 
Jones, Kirkby, & Brady [see Kempf, 1986: 
65] as is explicitly told by the ‘Table of 
Contents’ published with part 2 of that 
monograph in 1884, and not just Jones & 
Kirkby, as cited by many authors who per- 
haps did not know of the complete mono- 
graph.” 

Kornicker & Sohn (2000:20) had. fol- 


lowed Sylvester-Bradley (1951:210; 1961: 
Q403) in attributing C. oblonga to Jones & 
Kirkby 1874, not Jones, Kirkby, & Brady 
1874. The following discussion attempts to 
resolve which set of authors is correct. Be- 
cause the Jones & Kirkby 1874 publication 
includes an additional 35 new species and 
several new varieties, any conclusion drawn 
herein concerning proper attribution of the 
junior homonym Cypridina oblonga should 
also apply to them (no attempt is made 
herein to determine whether or not some of 
the additional species are junior homonyms 
or junior synonyms). 

For brevity, the 1874 and 1884 volumes 
of the Palaeontographical Society are sim- 
ply referred to as “‘the 1874 volume” and 
“the 1884 volume’’, respectively, through- 
out this paper. 


Discussion 


Fig. | is a copy of the title page of the 
1874 volume of the Palaeontographical So- 
ciety. It clearly states that ‘Part I, The Cy- 
pridinadae and their allies’’ containing pag- 


VOLUME 114, NUMBER 3 


687 


A MONOGRAPH 


OY THE 


BRITISH FOSSIL 


BIVALVED ENTOMOSTRACA 


FROM THE 


CARBONIFEROUS 


FORMATIONS. 


PROFESSOR T. RUPERT JONES, F.R.S., G.S., &c. &e.; 
JAMES W. KIRKBY, ESQ., &c. &c.; 


GEORGE §. BRADY, ESQ., C.M.Z.S., &. &e. 


PART I. 
THE CYPRIDINADA AND THEIR ALLIES. 


By PROF. T. RUPERT JONES, F.B.S., G.S., axp J. W. KIRBY, Ese., dc. &c. 


CONTAINING 


Pacts 1—56; Pirates I—Y. 


LONDON: 
PRINTED FOR THE PALHONTOGRAPHICAL SOCIETY. 
1874. 


Fig. 1. 


es 1—56 and plates I—V, is the contribution 
of T. Rupert Jones & James W. Kirkby. On 
page 20 of that publication appears the de- 
scription of Cypridina oblonga having the 
heading “13. Cypridina oblonga. Sp. nov. 
Plate V, figs. 12 a—c.”’ 

The names Jones, Kirkby, & Brady are 
listed as authors of the whole monograph 
on the title pages of both the 1874 (Fig. 1) 
and 1884 (Fig. 2) publications of the Pa- 
laeontographical Society, but in comparison 


Title page of the 1874 volume of the Palaeontographical Society. 


with the 1874 title page (Fig. 1) where 
Jones & Kirkby are listed as the authors of 
Part 1, no authors are specified for Part 1, 
No. 2 in the 1884 title page. However, ““Di- 
rections to the binder” (Fig. 3) that presum- 
ably accompanied the distribution of the 
1884 volume instructed the binder to ““Can- 
cel the Title-pages in the Palaeontographi- 
cal volumes for the years 1874 [Fig. 1] and 
1884 [Fig. 2], and substitute the General Ti- 
tle page (dated 1874 and 1884) provided in 


688 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


A MONOGRAPH 


OF THE 


Syd A UIL Sloboda Que edie 


BIVALVED ENTOMOSTRACA 


FROM THE 


CARBONIFEROUS 


FORMATIONS. 


PROFESSOR T. RUPERT JONES, F.R.S., G.S., &e. &e.; 


JAMES W. KIRKBY, ESQ., &&. &.; 


PROFESSOR GHEORGHE §. BRADY, M.D., F.R.S., F.L.S., &c., &e. 


PART I. 


No. 2. 


THE CYPRIDINADA AND THEIR ALLIES. 


CONTAINING 


Pages i—i), 57—92; Prates VI, VII. 


With TitLE-pace anD Directions For BINDING. 


LONDON: 
PRINTED FOR THE PALHONTOGRAPHICAL SOCIETY. 
1884. 


Fig. 2. 


the volume for 1884” (Fig. 4). The page 
with directions to the binder also gives the 
dates of publication of the various parts of 
the bound volume (Fig. 3). In the General 
Title page (dated 1874 and 1884) Jones, 
Kirkby, & Brady are listed as authors of the 
whole monograph, but no authors are listed 
under Part 1 (Fig. 4). Unlike the 1874 vol- 


Title page of the 1884 volume_.of the Palaeontographical Society. 


ume, the 1884 volume contains a Table of 
Contents (pages i—i11). In that table Cypri- 
dina oblonga and 35 additional species des- 
ignated as new species in the 1874 volume 
are attributed to Jones, Kirkby, & Brady. 
Only one of the new species, Cypridina 
Bradyana (sic), is attributed to Jones & 
Kirkby. In 1874 (page 15) Jones & Kirkby 


VOLUME 114, NUMBER 3 


689 


THE CARBONIFEROUS BIVALVED ENTOMOSTRACA. 


PART I. 


DIRECTIONS TO 


THE BINDER. 


The Monograph of the Carboniferous Bivalved Entomostraca, Part I (the Cypridinadz and their 
Allies), will be found in the volumes of the Palzontographical Society for the years 1874 and 1884. —_ 


= = A 
ancel the Title-pages (dated 1874 and 1884) in the-purts issued in the Palseontographical-volumes 
for/the years 1874 and 1884 substitute une Grea eee 1874—1884) vided in the 


volume for T884. 


ORDER OF BINDING AND DATES OF PUBLICATION. 


PAGES | PLATES SOS a0 aL PUBLISHED 
Title-page — 1884 December, 1884 

1—ili —_— 

1—56 I—V 1874 July, 1874 

57—92 VI—VII 18&4 December, 1884 


Fig. 3. “Directions to the Binder” in the 1884 volume of the Palaeontographical Society. 


had explained the naming of that species as 
follows: “it is named after our accom- 
plished friend, Mr. G. S. Brady of Sunder- 
land, who has favoured us with much help 
in the study of these and other fossil En- 
tomostraca.”’ One species, Bradycynetus 
Rankinianus (sic), which is attributed to 
Jones & Kirkby 1867 in the 1874 volume, 
is attributed to Jones, Kirkby, & Brady in 
the Table of Contents of the 1884 volume. 
Another species, Cypridina radiata, which 
is attributed to Jones, Kirkby & Brady in 
the Table of Contents of the 1884 volume, 
is attributed to Jones & Kirkby in an “‘Ad- 
dendum to the Cypridinadae”’ in the 1884 
volume (page 81). The Table of Contents 
does not indicate which part of the volume 
was published in 1874, and which part in 
1884. 

Jones & Kirkby (1886:508, 509: Table 
II), in a table entitled, ‘““The Genera and 
Species of Carboniferous Ostracoda, and 
their Occurrences in England, Scotland, and 
Ireland” attributed Cypridina oblonga to 


Jones, Kirkby & Brady. Other species in the 
table that are also present in the Table of 
Contents in the 1884 volume are attributed 
to the same authors in both places. This in- 
dicates that Jones and Kirkby agreed in 
1886 with the addition of Brady’s name to 
C. oblonga in the 1884 volume. 

The number of 1874 volumes that have 
not been bound to an 1884 volume appears 
to be sparse. The shared on-line catalog da- 
tabase, Online Computer Library Center, 
Inc. (OCLC), of January, 2000, lists only 
the University of Oxford as having an 1874 
volume not bound to the 1884 volume. That 
1874 volume, which is in the Oxford Uni- 
versity Museum (Natural History) Library, 
has a title page identical to that of Fig. 1, 
which is a copy of the tithe page in an 
1874—1884 bound volume in the Smithson- 
ian Library (Parker, in litt., 4 Oct 2000). 
The 1874 volume at Oxford does not have 
a Table of Contents (Parker, in litt., 4 Oct 
2000). 

An abstract published by Jones (1873b: 


690 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


A MONOGRAPH 


OF THE 


Qjerrrcee mMOSert 


BIVALVED ENTOMOSTRACA 


FROM THE 


CARBONIFEROUS FORMATIONS. 


PROFESSOR T. RUPERT JONES, F.R.S., G.S., &c. &.; 


JAMES W. KIRKBY, ESQ., &e. &c.; 


PROFESSOR GHORGE Ss Sn AW Ye sep Wakes. eb las sccumcrc: 


WaQee JL 


THE CYPRIDINADA AND THEIR ALLIES. 


LONDON : 
PRINTED FOR THE PALHZONTOGRAPHICAL SOCIETY. 
1874—1884. 


Fig. 4. 
Palaeontographical Society. 


409) mentions Cypridina oblonga (page 
410) and other new species that subsequent- 
ly were described in the 1874 and 1884 vol- 
umes. A description of C. oblonga is not 
given; therefore, the species name is a no- 
men nudum. The abstract states (page 409): 


Substitute “General Title-page (dated 1874-1884)” of the combined 1874-1884 volumes of the 


‘‘Numerous other specimens from Ireland, 
Scotland, Yorkshire, Derbyshire, and the 
Isle of Man, communicated by friends, have 
also been studied; and the results will be 
given in detail in a Monograph by Messrs. 
Jones and Kirkby, to be published by the 


VOLUME 114, NUMBER 3 


691 


Table 1.—List of selected authors (arranged chronologically) showing attribution by them of authorship of 
five genera of Paleozoic Ostracoda. (B = George S. Brady, J. = T. Rupert Jones, K = James W. Kirkby, — = 
genus not mentioned; list does not include Jones, 1873a, b.) 


Attribution 

Author Cypridinella Cypridellina Sulcuna Rhombina Offa 
Jones & Kirkby (1874) J&K J&K JI&K J&K J&K 
Jones, Kirkby, & Brady (1884) I,K &B I,K &B I,K &B J,K&B I,K&B 
Woodward (1877) J&K J&K I,K &B IJK &B K&B 
Jones & Kirkby (1887) J J J J J 
Gemmellaro (1890) J&K JI&K — — = 
Bassler & Kellett (1934) I,K &B J,K&B I,K &B I,K&B JI,K&B 
Neave (1939) I,K &B I,K &B == == a 
Neave (1940a) — — — — J,K&B 
Neave (1940b) — — J J,K&B — 
Sylvester-Bradley (1951) — — — J&K — 
Howe (1955) J J&K J J J 
Mertens (1958) J J&K J J&K J 
Pokorny (1958, 1965) J J — I,K &B J 
Sylvester-Bradley (1961) J&K J&K J&K J&K J&K 
Howe (1962) J J&K J&K J&K J&K 
Kempf (1986, pt. 1) I,K &B J,K&B I,K &B I,K &B IJK&B 
Kormicker & Sohn (2000) J&K J&K J&K — — 
Herein J J J J J 


Palaeontographical Society, and now in 
press.” The only importance of the abstract 
in the Quarterly Journal (Jones, 1873b) to 
the present study is that it contains a ref- 
erence to the monograph by Jones & Kirk- 
by then in press by the Palaeontographical 
Society. 

Concerning the Palaeontographical So- 
ciety archives, Paul Ensom (in litt., 12 Dec 
2000) kindly sent the following: 

“I checked all the Annual Reports (print- 
ed), which were present in the minute book, 
1.e., tabbed in. There was no reference to 
the problem [addition of Brady’s name]. 
The Printed annual reports only appear in 
1875, 1976, then 1879 onwards; 1877 and 
1878 are not present. The annual report for 
1884 states “The other portions of the Vol- 
ume for 1884 will be. . the continuation of 
the Carboniferous Entomostraca by Messrs 
Prof. Jones, J.W. Kirkby, and Prof G.S. Bra- 
dy,..., and that is all.” 

While researching the above, I observed 
that five Paleozoic ostracode genera (Cy- 
pridinella, Cypridellina, Sulcana, Rhombi- 
na, and Offa), which had been described by 


Jones (1873a), were designated as new gen- 
era in both Jones & Kirkby (in Jones, Kirk- 
by, & Brady 1874) and Jones, Kirkby, & 
Brady 1884. Since then various authors 
have attributed the genera to either Jones 
1873, or Jones & Kirkby (in Jones, Kirkby, 
& Brady 1874), or Jones, Kirkby, & Brady 
1884 (Table 1). The descriptions of the five 
genera in Jones 1873a are adequate to sat- 
isfy Article 12.1 of the International Code 
of Zoological Nomenclature, Fourth Edi- 
tion (International Commission on Zoolog- 
ical Nomenclature 1999). Therefore, the 
genera should be attributed to Jones. 


Conclusions 


The 1874 volume satisfies the “Criteria 
of Publication” (Chapter 3) and “Criteria 
of Availability’? (Chapter 4) of the Inter- 
national Code of Zoological Nomenclature, 
Fourth Edition. The Abstract published by 
Jones (1873b:409) indicates that only Jones 
& Kirkby were intended to be authors of 
Part 1 of the 1874 volume, and the original 
title page of the 1874 volume should be ac- 


692 


cepted at face value, not withstanding con- 
trary information issued 10 years later. 

My interpretation of rules listed under 
“Date of Publication’? (Chapter 5) in the 
Code and other provisions contained therein 
leads me to believe that only Jones and 
Kirkby are the authors of the junior hom- 
onym Cypridina oblonga in the 1874 vol- 
ume. 


Acknowledgments 


I wish to thank Eugen Karl Kempf for 
making me aware of this problem and for 
correspondence concerning it, including a 
suggestion for the new specific name. I also 
thank Andrew Richard Parker, Oxford Uni- 
versity, for information mentioned above, 
and Elizabeth Harrison-Nelson, Smuithsoni- 
an Institution, for assistance in preparation 
of the paper. Paul C. Ensom, John E. Whit- 
taker, and C. Giles Miller, The Museum of 
Natural History, London, kindly answered 
correspondence concerning the 1874—1884 
volumes, and Wendy Cawthorne, Geologi- 
cal Society Library, London, kindly an- 
swered correspondence concerning the 
Jones (1873b) abstract. I thank Frederic M. 
Swain, University of Minnesota, and sev- 
eral anonymous reviewers for criticizing the 
manuscript. I much appreciate the review of 
the manuscript by Brian Kensley (Smith- 
sonian Institution). I also thank for their 
help Leslie Overstreet, David Steere, and 
Martha Rosen, Smithsonian Institution; 
Storrs Olson, Smithsonian Institution; and 
F Christian Thompson, U.S. Department of 
Agriculture. 


Literature Cited 


Bassler, Ray S. & Betty Kellett. 1934. Bibliographic 
index of Paleozoic Ostracoda.—Special Papers 
of the Geological Society of America 1. Geo- 
logical Society of America, New York, 500 pp. 

Gemmellaro, G. 1890. II crostacei dei calcari con fu- 
sulina della valle del fiume.—Rendiconti. Ac- 
cademia nazionale dei Lincei (Rome) series 3, 
8(1):1—40. 

Grube, E. 1859. Bemerkungen iiber Cypridina und 
eine neue Art dieser Gattung (Cypridina oblon- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ga).—Archiv fiir Naturgeschichte (Berlin) 25: 
322-337. 

Howe, H. V. W. 1955. Handbook of ostracod taxono- 
my. Louisiana State University Studies (Physi- 
cal Science Series) 1, Louisiana State Univer- 
sity Press, Baton Rouge, 386 pp. 

. 1962. Ostracod taxonomy. Baton Rouge, Lou- 
isiana State University Press, 366 pp. 

International Commission on Zoological Nomencla- 
ture. 1999. International Code of Zoological 
Nomenclature, Fourth Edition. The Internation- 
al Trust for Zoological Nomenclature, London, 
306 pp. 

Jones, T. R. 1873a. On ancient water-fleas of the Os- 
tracodous and Phyllopodus tribes (Bivalved En- 
tomostraca). Part Il—The Cypridinidae.—The 
Monthly Microscopical Journal of the Royal 
Microscopical Society 10:71—78. 

Jones, T. R. 1873b. On some bivalved Entomostraca, 
chiefly Cypridinidae, of the Carboniferous For- 
mations.—Quarterly Journal of the Geological 
Society of London 29:409—412. 

, & J. W. Kirkby. 1867. On the Entomostraca 

of the Carboniferous rocks of Scotland. 

Transactions of the Geological Society of 

Glasgow 2(3):213—228. 

» & . 1874. Part 1. The Cypridinadae 

and their allies. Im T. R. Jones, J. W. Kirkby, & 

G. S. Brady, A monograph of the British fossil 

bivalved Entomostraca from the Carboniferous 

Formations. Palaeontographical Society (Lon- 

don):56 pp. 

, & . 1886. Notes on the distribution of 

the Ostracoda of the Carboniferous Formations 

of the British Isles —The Quarterly Journal of 

the Geological Society of London 42:496—514. 

, & . 1887. A list of the genera and 

species of bivalved entomostraca found in the 

Carboniferous formations of Great Britain and 

Ireland, with notes on the genera and their dis- 

tribution.—Proceedings of the Geologists’ As- 

sociation 9(7):495—515. 

, & G. S. Brady 1884. A monograph 
of the British fossil bivalved Entomostraca from 
the Carboniferous Formations. Part I. No. 2. 
The Cypridinadae and their allies —Palaeonto- 
graphical Society (London):57—92. 

Kempf, E. K. 1986. Index and bibliography of marine 
Ostracoda 2, Index C.—Geologisches Institut 
der Universitaet zu Koeln Sonderveroeffentli- 
chungen 52:1—774. 

Kornicker, L. S., & I. G. Sohn. 2000. Myodocopid 
Ostracoda from the Late Permian of Greece and 
a basic classification for Paleozoic and Meso- 
zoic Myodocopida.—Smithsonian Contribu- 
tions to Paleobiology 91:1—33. 

Mertens, E. 1958. Zur Kenntnis der Ordnung Ostra- 


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coda (Crustacea).—Geologische Jahrbuch 
(Hannover) 75:311—318. 

Neave, S. A. 1939. Nomenclator Zoologicus 1 (A-C). 
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957 pp. 

. 1940a. Nomenclator Zoologicus 3 (M—P). The 

Zoological Society of London, London, 1065 

Pp. 

. 1940b. Nomenclator Zoologicus 4 (Q—Z). The 
Zoological Society of London, London, 758 pp. 

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ary 2000. 6565 Frantz Road, Dublin, Ohio 
43017-3395, U.S.A. (http://purl.org/oclc/pi) 

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Wissenschaften, Berlin, 453 pp. 


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. 1965. Principles of zoological micropalaeon- 
tology 2. Pergamon Press, Oxford, 465 pp. 
[Translated by K. A. Allen, edited by J. W. Nea- 
le]. 

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new family of Carboniferous Ostracoda.—Geo- 
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. 1961. Myodocopida. Pp. 387—406 in Treatise 
on Invertebrate Paleontology—Part Q, Arthrop- 
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Woodward, H. W. 1877. Catalogue of British fossil 
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
114(3):694—707. 2001. 


Seepiophila jonesi, a new genus and species of vestimentiferan tube 
worm (Annelida: Pogonophora) from hydrocarbon seep communities 
in the Gulf of Mexico 


Stephen L. Gardiner, Erin McMullin, and Charles R. Fisher 


(SLG) Department of Biology, Bryn Mawr College, 101 N. Merion Avenue, Bryn Mawr, 
Pennsylvania 19010, U.S.A.; 
(EM, CRF) The Pennsylvania State University, Department of Biology, 208 Mueller Laboratory, 
University Park, Pennsylvania 16802, U.S.A. 


Abstract.—Seepiophila jonesi, a new genus and species of vestimentiferan 
tube worm is described from material collected from hydrocarbon seep com- 
munities in the Gulf of Mexico, based on morphological and molecular char- 
acters. In these communities, individuals occur singly, in small groups or in 
association with large aggregations of a second vestimentiferan species, La- 
mellibrachia cf. luymesi. Tubes extend deeply into the sediment, and a rela- 
tively short apical portion normally projects above the sea floor. Seepiophila 
jonesi is most similar to vestimentiferan species in the genus Escarpia but 
differs from it and other vestimentiferan species by a combination of morpho- 
logical characters that includes the number of types of branchial filaments as- 
sociated with the branchial plume, the presence of an incision in the poster- 
oventral margin of the vestimentum, the presence of a well-developed, medial 
obturacular structure, the relative position of the excretory pore and the pres- 
ence of a variable number of broad collars associated with the tube. The di- 
vergence in the sequence of the cytochrome oxidase I gene between S. jonesi 
and other described vestimentiferan tube worms supports its placement in a 
new genus within the family Escarpiidae. 


With the description of Lamellibrachia 
barhami, Webb (1969) recorded the first 
vestimentiferan tube worm from a cold-wa- 
ter site in the northeastern Pacific Ocean. 
The discovery of hydrothermal vents and 
their associated fauna along the Galapagos 
Rift, East Pacific Rise and Juan de Fuca 
Ridge of the eastern Pacific Ocean and in 
various regions of the western Pacific has 
led to the description of ten additional ves- 
timentiferan species from the Pacific Ocean 
(Jones 1981, 1985; Southward 1991; Miura 
et al. 1997; Southward & Galkin 1997; 
Southward et al. 2001). 

In contrast to the Pacific Ocean, the At- 
lantic Ocean, including the Gulf of Mexico, 
at present, exhibits much less vestimenti- 
feran diversity. Two species of Lamelli- 


brachia are known from putative cold-wa- 
ter sites in the western Atlantic Ocean. Van 
der Land & N@rrevang (1975) described L. 
luymesi based on a single male specimen 
taken from about 500 m of water off Guy- 
ana, and L. victori was characterized by 
Mané-Garzon & Montero (1985) based on 
several specimens dredged from about 300 
m of water on the continental slope of Uru- 
guay. 

Paull et al. (1984) were the first to report 
the presence of vestimentiferan tube worms 
in the Gulf of Mexico. Diving in the sub- 
mersible Alvin at a depth of more than 3000 
m at the base of the Florida Escarpment, 
they observed clumps of specimens that 
varied in density from a few to more than 
100 individuals per square meter. Speci- 


VOLUME 114, NUMBER 3 


695 


Table 1.—Names, coordinates and depths of hydrocarbon seep sites on the Louisiana Slope, Gulf of Mexico 


sampled for the present study. 


Lease block Site name Latitude Longitude Depth (m) 
GC185 Bush Hill 27°47'N SiS) WY 540 
GC233 Brine Pool 27°43.4'N 91°16.8"W 640 
GC234 Green Canyon 27°44.1'N 91°15.3'W 540 
TAMU-17 Tamu 27°43.8'N 91°18.2'W 560 
GB425 Garden Banks 2] 333) N 92°32.4'W 570 


mens collected from this site were later giv- 
en the name of Escarpia laminata by Jones 
(1985). 

As part of an investigation of the possible 
effects of hydrocarbon release into overly- 
ing waters in the Gulf of Mexico, Kennicutt 
et al. (1985) performed deep-water trawls 
at two sites along the Louisiana Slope. One 
of the trawls contained vestimentiferan 
specimens which they identified as Lamel- 
librachia sp. Subsequent cruises and sub- 
mersible dives to sites along the Louisiana 
Slope revealed the presence of dense as- 
semblages of hydrothermal vent-type taxa, 
including large aggregations of the vesti- 
mentiferan Lamellibrachia cf. luymesi (see 
MacDonald et al. 1989, among others). Oc- 
curring in association with these aggrega- 
tions, or sometimes occurring independent- 
ly, is a second undescribed vestimentiferan 
species (CRE SLG, EM pers. obv.). The 
purpose of the present study is to provide a 
description of this new species, using mor- 
phological features, and to examine its re- 
lationships with other vestimentiferans, us- 
ing morphological and molecular analyses. 


Materials and Methods 


Specimens were collected by the manned 
submersibles Johnson Sea Link I (JSL I) 
and Johnson Sea Link IT (JSL I) from five 
seep sites on the Louisiana Slope in the 
Gulf of Mexico, ranging from 540 to 640 
m in depth: Bush Hill, GC234, Brine Pool, 
TAMU-17 and GB425 (Table 1). Speci- 
mens were brought to the surface in a tem- 
perature-insulated container and immediate- 
ly placed in chilled water on board ship. 


Animals collected for morphological stud- 
ies were removed from their tubes, pre- 
served in 10% buffered formalin in sea- 
water and subsequently transferred to 70% 
ethanol. Specimens collected for molecular 
analyses were removed from their tubes, 
and a sample of the vestimentum was 
snipped and immediately frozen in liquid 
nitrogen. Frozen samples were transported 
on dry ice and subsequently stored at 
=ONC; 

To show the relationship of the Louisiana 
Slope vestimentiferans to other known ves- 
timentiferans, we sequenced a portion of 
the mitochondrial cytochrome oxidase I 
gene (COI) and compared it with published 
data for other vestimentiferans (Black et al. 
1997, Kojima et al. 1997, Feldman et al. 
1998). The four specimens of the new spe- 
cies described herein were collected from 
four different sites within 65 km of each 
other on the Louisiana Slope (Bush Hill, 
GC234, TAMU-17 and GB425). Specimens 
were stored at —80°C before and after DNA 
extraction. For these studies, total nucleic 
acids were extracted from vestimentum tis- 
sue by the classic phenol/chloroform ex- 
traction technique. The COI gene was am- 
plified by PCR, at an annealing temperature 
of 55°C, with the following primers: COIf: 
TC(CA)ACTAATCA(CT) AA(GA) GA- 
(CT)ATTGG(ATGC)AC, COIr:CC(ATG)- 
CTTAG(TA)CCTA(GA)(GA)AA (GA)TG- 
TTG(ATCG)GG (Nelson & Fisher 2000). 
PCR products were visualized on a 1.5% 
agarose gel (GIBCO BRL) stained with eth- 
idium bromide. Products were cleaned with 
a PCR Purification Kit (QIAGEN) before 


696 


sequencing. This primer pair amplifies an 
approximately 1250 base pair segment, 
from which a 1100 base pair fragment was 
sequenced with multiple primers, generat- 
ing overlapping sequences from both 
strands. 

Cycle sequencing PCR was done with a 
Beckman CEQ DTCS dye terminator re- 
action kit (BECKMAN), under manufactur- 
er’s suggested conditions. End labeled prod- 
ucts were separated on a capillary autose- 
quencer (Beckman CEQ 2000XL). Individ- 
ual sequence runs were assembled and 
edited with SEQMAN (DNASTAR, Inc.). 
Consensus sequences generated in this 
study were aligned (MEGALIGN, DNA- 
STAR Inc.) with those generated in previ- 
ous studies available in Genbank (Table 2). 
Molecular evolutionary relationships 
among sequences were examined by the 
minimum evolution (ME) method of tree 
construction (Rzhetsky & Nei 1992) based 
on pairwise genetic distances estimated 
from the proportion of differences and cor- 
rected for multiple substitutions by the Ki- 
mura 2-parameter formula (Kimura 1980). 
Significance of branching order was eval- 
uated by bootstrap analysis with 1000 rep- 
lications. Both a pogonophoran from the 
study of vestimentiferans in Japan by Ko- 
jima et al. (1997) and Galathealinum bra- 
chiosum, a described pogonophoran spe- 
cies, were used as outgroup sequences. All 
analyses were performed with MEGA (Ku- 
mar, version 2.0 beta). 


Systematics 


The systematic relationships of vestimen- 
tiferans (= subphylum Obturata sensu 
Jones 1981) to pogonophorans (= subphy- 
lum Perviata sensu Jones 1981) and to other 
invertebrate taxa such as the Annelida, as 
well as their placement in a taxonomic hi- 
erarchy, remain unsettled. It is not a pur- 
pose of the present study to provide an ex- 
haustive historical account of these issues. 
Recent morphological, molecular and de- 
velopmental studies suggest a close rela- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


tionship between vestimentiferans and po- 
gonophorans and also support an annelid 
affinity for both groups or place them with- 
in the Annelida or Polychaeta (Southward 
1988, 1999; Gardiner & Jones 1994; Rouse 
& Fauchald 1995, 1997; Black et al. 1997; 
Kojima et al. 1997; McHugh 1997, 2000; 
Bartolomaeus 1998; Halanych et al. 1998; 
Rouse 2001). In spite of the seemingly con- 
vincing body of evidence provided by these 
studies, Salvini-Plawen (2000) questioned 
the reliability of characters used in deter- 
mining possible relationships between ves- 
timentiferans and pogonophorans and the 
Annelida-Polychaeta. He concluded that the 
current level of knowledge of characters of 
vestimentiferans and pogonophorans is in- 
sufficient to provide definitive relationships. 
Rather, he suggested that, depending on 
which characters are being analyzed, the 
two groups demonstrate possible affinities 
to the Annelida-Polychaeta, Annelida-Ow- 
eniidae and Oligomera. He stated that ad- 
ditional comprehensive studies of vestimen- 
tiferans and pogonophorans are required to 
refine better our understanding of their pos- 
sible relationships with other taxa. 

In order to maintain consistency with 
other recent descriptions of vestimentifer- 
ans (Southward 1991, Southward et al. 
1995, Miura et al. 1997, Southward & 
Galkin 1997, Southward et al. 2001), we 
here adopt the classification proposed by 
Southward (1991) and consider vestimen- 
tiferans as comprising a subclass within the 
class Pogonophora of the phylum Annelida. 


Subclass Obturata Jones, 1981 
Order Basibranchia Jones, 1981 
Family Escarpiidae Jones, 1985, amended 


Diagnosis.—Vestimentiferan worms with 
tapering tubes and bodies. Anterior obtu- 
racular region provided with branchial 
plume; orientation of branchial lamellae, 
relative to obturaculum, axial and parallel; 
branchial filaments of obturacular plume of 
one type or grouped in two types; plume 
lacking peripheral lamellar sheaths; anterior 


VOLUME 114, NUMBER 3 


face of obturaculum with moderate to thick 
crust and usually with variously developed 
medial structure arising from between ob- 
turacular halves; paired internal excretory 
ducts opening by single, dorsal medial ex- 
cretory pore near base of obturaculum or 
more anteriorly, up to one-third distance of 
obturaculum from base. Anteroventral mar- 
gin of vestimentum incised, posteroventral 
margin entire or incised. Opisthosome with 
setae in single to triple rows. Tube hard, 
tapering, with or without anterior funnel 
and variable number of broadly flared ex- 
ternal collars and with or without variable 
number of rings along length. 

Remarks.—The family Escarpiidae was 
amended by Southward et al. (2001) to al- 
low for the inclusion of a new genus and 
species collected from deep water in the 
western Pacific Ocean. 

The amended diagnosis of the family Es- 
carpiidae provided above is constructed 
from observations by one of us (SLG) of 
type specimens of Escarpia spicata and E. 
laminata and the new genus and species de- 
scribed herein, together with characters of 
the new genus described in Southward et 
al. (2001). The significant morphological 
feature included in this amended diagnosis 
is the presence of only a single type of 
branchial filament associated with the ob- 
turacular plume. 

Jones (1985) did not possess complete 
specimens of Escarpia spicata or E. lami- 
nata. Therefore, he was unable to charac- 
terize the opisthosome of Escarpia. South- 
ward (pers. comm., Southward et al. 2001) 
describes the opisthosome of one specimen 
of their new genus as comprising about 15 
segments with setae in single to triple rows. 
Setae are composed of an anterior smaller 
group of 2—3 teeth and a larger posterior 
group of teeth occurring in 3—4 rows. 


Seepiophila, new genus 


Diagnosis.—Vestimentiferan worms with 
tapering tubes and bodies. Anterior obtu- 
racular region provided with branchial 


697 


plume; orientation of branchial lamellae, 
relative to obturaculum, axial and parallel; 
branchial filaments of obturacular plume of 
one type; plume lacking peripheral lamellar 
sheaths; anterior face of obturaculum with 
moderate crust and with conspicuous me- 
dial structure arising from between obtura- 
cular halves; paired internal excretory ducts 
opening by single, dorsal medial excretory 
pore situated on obturaculum approximate- 
ly one-third distance anteriorly from base. 
Anteroventral and posteroventral margins 
of vestimentum incised. Opisthosome un- 
known. Tube hard, tapering, aperture sur- 
rounded by broadly flaring funnel, with var- 
iable number of broadly flared external col- 
lars anteriorly and with variable number of 
rings along length. 

Type species.—Seepiophila jonesi, new 
species, by present designation. 

Gender.—Feminine. 

Etymology.—From English seep (= a 
place where water or petroleum oozes out 
slowly) + Greek philia (= affection, fond- 
ness), in reference to the strong preference 
of these worms to inhabit hydrocarbon seep 
communities in the deep sea. 


Seepiophila jonesi, new species 
Figs. 1—5 


Unidentified vestimentiferan.—Childress et 
aleelOsoalsOre 

Undescribed family.—Brooks et al., 1987: 
1139, table 1. 

Escarpia-like [sic].—Brooks et al., 1987: 
1140, table 2, 1141, table 3. 

Undescribed member of the family Escar- 
piidae.—Fisher et al., 1988:230, 232. 

Escarpia-like.—MacDonald et al., 1989: 
239, 243, table 4.—Childress & Fisher, 
MO ODES SP 

The escarpiid.—MacDonald et al., 1989: 
241, 245, fig. 3B. 

Escarpia sp. {not Escarpia Jones, 1985].— 
MacDonald et al., 1989:235.—Young et 
al., 1996:514—516, figs. 1b, 2a, c, f-1.— 
Miura et al., 1997:455.—Southward, 
1999:196.—Tyler & Young, 1999:197, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 
crust-like material (cr) and prominent medial process (asterisk). bf, branchial filaments. B, Enlargement of portion 
of medial process. Note numerous spines along lateral margins. Scale bars: A = 2 mm; B = | mm. 


table 3, 198.—Salvini-Plawen, 2000:133, 
134. 

Undescribed genus.—Fisher, 1990:406, ta- 
ble 1, 410. 

Unidentified escarpid.—Fisher et al., 1990: 
1095, table 1. 

E. sp. [not Escarpia Jones, 1985].—Fisher, 
1995:307, table 4. 

Escarpiidae species.—Nelson & Fisher, 
1995:128, table 1. 

Escarpia spicata {not Escarpia spicata 
Jones, 1985].—Feldman et al., 1997:270. 

Escarpid-like.—Fisher et al., 1997:83—94, 
fig. 1b (as escarpids).—Nelson & Fisher, 
2000: 1—10. 

Undescribed seep escarpid.—Scott et al., 
1998:379, table 1. 


Type material.—See Table 1 for coordi- 
nates and depths of hydrocarbon seep com- 
munities. Holotype, (USNM 188519), 6 
paratypes (USNM 188520—188525), Bush 
Hill hydrocarbon seep community, JSL J 
Dive 3775, 8 Oct 1994, coll. S. L. Gardiner; 
6 paratypes, Bush Hill hydrocarbon seep 
community, JSL J Dive 3530, 28 Jun 1993, 
coll. J. J. Childress. Additional type mate- 
rial is deposited in the Natural History Mu- 
seum, London, the Canadian Museum of 
Nature, the National Science Museum, To- 
kyo and the Muséum National d’ Histoire 
Naturelle, Paris. 


Seepiophila jonesi, new species. Holotype (USNM 188519). A, Anterior face of obturaculum showing 


Additional material examined.—See Ta- 
ble 1 for coordinates and depths of hydro- 
carbon seep communities. For morpholog- 
ical analyses: 14 specimens, Green Canyon 
hydrocarbon seep community, JSL J Dive 
3525, 26 Jun 1993, coll. BE. NixeDeseeer 
mens, Green Canyon hydrocarbon seep 
community, JSL J Dive 3526, 26 Jun 1993, 
coll. J. J. Childress; 8 specimens, Bush Hill 
hydrocarbon seep community, JSL I Dive 
3530, 28 Jun 1993, coll. J. J. Childress; 17 
specimens, Bush Hill hydrocarbon seep 
community, JSL I Dive 3775, 8 Oct 1994, 
coll. S. L. Gardiner. For molecular analyses: 
1 specimen, Bush Hill hydrocarbon seep 
community, JSL IT Dive 2853, 10 Jul 1997, 
coll. K. Nelson; 1 specimen, Green Canyon 
hydrocarbon seep community, JSL IJ Dive 
2866, 16 Jul 1997, coll. K. Nelson; 1 spec- 
imen, Garden Banks hydrocarbon seep 
community, JSL J Dive 4053, 17 Jul 1998, 
coll. E. McMullin; 1 specimen, TAMU-17 
hydrocarbon seep community, JSL I Dive 
4049, 15 Jul 1998, coll. E. McMullin. 

Diagnosis.—Characters of the genus. 

Etymology.—The species is named in 
honor of the late Meredith L. Jones whose 
studies of vestimentiferan anatomy, mor- 
phology and development contributed 
greatly to our understanding of this enig- 
matic group of marine worms. 


VOLUME 114, NUMBER 3 


Description.—Measurements of selected 
specimens (length by diameter, in mm; ob- 
turacular: vestimental: trunk: opisthosomal 
regions; + = incomplete; — = change in 
diameter; — = missing); (USNM 188519): 
9.5 by 7.5: 36.4 by 10.2: 415+ by 5.9-0.4: 
—; (USNM 188520): 12.1 by 7.8: 44.2 by 
8.9: 330+ by 6.4-1.6: —; (USNM 
188521): 11.5 by 6.1: 38.7 by 7.5: 100+ by 
4.5—3.5: —; (USNM 188522): 10.7 by 6.1: 
45.1 by 6.5: 380+ by 6.4—0.8: —; (USNM 
188525): 11.2 by 5.6: 52.1 by 7.4: 350+ by 
4.7-1.0: —). Measurements of selected 
tubes: (USNM 188519): 870+ by 12.4—1.0; 
(USNM 188520): 955+ by 9.6—0.8; 
(USNM_ 188523): 810+ by 11.6—0.6; 
(USNM 188524): 820+ by 10.9-2.7. 

Anterior face of obturaculum with mod- 
erately developed crust, up to approximate- 
ly 1 mm in thickness (Fig. 1A); cuticular 
axial rod between obturacular halves of dis- 
tal one-third of obturaculum, extending be- 
yond anterior face of obturaculum as prom- 
inent medial structure, laterally compressed, 
terminating bluntly, lateral margins provid- 
ed with numerous small spines (Fig. 1A, 
B); measurements of selected medial struc- 
tures (anteroposterior length above anterior 
face of obturaculum by dorsoventral height 
by lateral width) in mm: (USNM 188519): 
1.5 by 5.9 by 3; (USNM 188521): 1.6 by 
4.7 by 2.3; (USNM 188523): 1.5 by 5.5 by 
2.5; (USNM 188525): 0.8 by 4.5 by 1.7. 
Obturaculum with approximately 45—50 
pairs of branchial lamellae (Fig. 2A—D), 
lacking specialized sensory filaments; fila- 
ments comprising lamellae of one type, 
fused for greater portion of length, with sin- 
gle row of pinnules extending along free 
portion of filaments (Fig. 3A) and with two 
rows of ciliated cells (Fig. 3A, B); pinnules 
on filaments of lamella one (= oldest la- 
mella) relatively inconspicuous, often irreg- 
ularly spaced along distal length of fila- 
ments (Fig. 3C); pinnules on filaments of 
middle and posterior-most lamellae larger, 
regularly spaced along distal length of fil- 
aments (Fig. 3A, B); midventral face and 
distal sides of obturaculum bare; obturacu- 


699 


lum laterally compressed at base, oval to 
somewhat spindle-shaped in cross-section 
more distally (Fig. 4A), lacking dorsal 
groove, with poorly developed ventral 
ridge; paired internal excretory ducts (Fig. 
4A) opening by single pore on dorsal me- 
dial ridge approximately one-third distance 
of obturaculum from base (Fig. 2D, arrow), 
dorsal medial ridge sometimes darkly pig- 
mented; ratio of obturacular length to ves- 
timental length variable, 1:2.8 to 1:6.6. An- 
terior margin of vestimentum forming short 
sheath around base of obturaculum (Fig. 
2A), with shallow midventral incision often 
forming small lobes (Fig. 2C); posteroven- 
tral margin of vestimentum deeply incised, 
with two conspicuous lobes (Fig. 2C, large 
arrowhead, E); ratio of vestimental diameter 
to vestimental length highly variable, 1:3.1 
to 1:12.6; ventral surface with tear-drop 
shaped ciliated field; plaques associated 
with epidermis ventrolaterally (Fig. 2E, ar- 
rowhead); males with paired dorsal ciliated 
grooves extending from gonopores to near 
anterior end of vestimentum, converging 
slightly anteriorly (Fig. 2D, arrowheads). 
Trunk very long in adult specimens, taper- 
ing to less than 1 mm in diameter; epider- 
mis with numerous associated plaques (Fig. 
2E, arrowhead). Opisthosome unknown. 
Tube hard, often somewhat darkly colored, 
tapering posteriorly, aperture with broadly 
flaring funnel (Figs. 4B, 5); diameter of ap- 
erture variable (6.6-11 mm; X = 9.1 + 1.3 
mm, 2 = 34), with variable number of col- 
lars posterior to aperture (0-10; X = 3.2 + 
1.9, n = 34); number of rings posterior to 
aperture highly variable (11-59; X = 29.8 
+ 10.2, n = 34). 

Distribution.—Presently known only in 
association with certain hydrocarbon seep 
communities in the Gulf of Mexico be- 
tween 500 and 1000 m depth. 

Remarks.—Seven vestimentiferan spe- 
cies in Six genera possess a medial structure 
in association with the obturaculum: Escar- 
pia spicata Jones, 1985, E. laminata Jones, 
1985, Oasisia alvinae Jones, 1985, Ridgeia 
piscesae Jones, 1985, Tevnia jerichonana 


700 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Seepiophila jonesi, new species. Holotype (USNM 188519). A, Dorsal view of obturacular (ob) and 
vestimental (v) regions and portion of trunk region (tr). Note broadly splayed anterior face of obturaculum. B, 
Left lateral view of obturacular region showing branchial filaments (bf), vestimental region with vestimental 
wings (vw) and portion of trunk. C, Ventral view of obturacular region, vestimental region with tear-drop shaped 
ciliated field (cf) and portion of trunk region. Small arrowheads indicate position of intraepithelial nerve cords. 
Note deeply incised posteroventral margin of vestimentum (large arrowhead). bf, branchial filaments. D, Enlarged 
dorsal view of obturacular region showing prominent dorsal ridge (dr) at base of obturaculum. Arrow indicates 


VOLUME 114, NUMBER 3 701 


Fig. 3. 


Seepiophila jonesi, new species. Scanning electron microscopy. Non-type specimens. A, Branchial 
filament from middle lamella. Single row of pinnules extends from near tip of filament to region where filament 
fuses with adjacent filaments. Note diminished size of pinnules in region of fusion (arrow). Arrowheads indicate 
cilia. B, Branchial filament from basal lamella. Note well-developed pinnules and two rows of ciliated cells (ci). 
C, Branchial filament from lamella one (= oldest lamella). Note single row of small, somewhat irregularly 


spaced pinnules. ci, cilia. Scale bars: A = 50 wm; B = 25 wm; C = 30 pm. 


Jones, 1985, a new genus described in 
Southward et al. (2001) and Seepiophila 
jJonesi, new species. Seepiophila jonesi, 
however, differs from all other species that 
possess an obturacular medial structure in 
having only a single type of filament com- 
prising the branchial lamellae. In S. jonesi, 
all filaments possess a single row of pin- 
nules, whereas other species that display an 
obturacular medial structure possess at least 
some filaments that lack pinnules. 

A crust-like material on the anterior face 
of the obturaculum and a medial structure 
secreted by the obturacular halves are char- 


acters shared by all species in the Escarpi- 
idae. The crust-like material appears to be 
about equally developed in the four species 
of the family. This is not the case, however, 
for the medial structure. Jones (1985) did 
not report dimensions for the medial struc- 
ture of Escarpia laminata, presumably be- 
cause of its small size. He reported, how- 
ever, that the medial structure of E. spicata 
extends as much as 9.5 mm above the an- 
terior face of the obturaculum. Southward 
(pers. comm.) indicates that one specimen 
of their new genus (Southward et al. 2001) 
possesses a medial structure that extends 20 


<< 


approximate position of excretory pore. Note ciliated groove on dorsal surface of vestimentum (arrowheads). E, 
Enlarged view of posteroventral margin of vestimentum showing lobed condition. Arrowheads indicate plaques 
associated with epidermis of vestimentum and trunk. cf, ciliated field. Scale bars: A-C = 10 mm; D = 5 mm; 
E = 2 mm. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Seepiophila jonesi, new species. A, Non-type specimen. Light microscopic view of transverse section 
through obturacular region. Small arrowheads indicate positions of paired excretory ducts. Arrows indicate 
sections through obturacular blood vessel. bf, branchial filaments; dr, dorsal ridge; ob, obturacular half. B, 
Anterior region of tube showing aperture surrounded by broadly flaring funnel and two well-developed collars 
posterior to aperture. Scale bars: A = 0.5 mm; B = 10 mm. 


Fig. 5. Individuals of Seepiophila jonesi (arrows) within a clump of juvenile specimens of Lamellibrachia 
cf. Juymesi and several seep mussels (m). No scale. 


VOLUME 114, NUMBER 3 


Escarpia sp. (Nankai Trough, 1200m) 


00 | Seepiophila jonesi (GC234, GoM, 550m) 


703 


Seepiophila jones! (GB425, GoM, 550m) 
Seepiophila jonesi (TAMU-17, GoM, 550m) 


Seepiophila jonesi (Bush Hill, GoM, 550m) 


oOoaQ--—-T AMA 0M 


A 400 'Escarpia sp. (Iheya Ridge, 1400m) 
Escarpia sp. (Nankai Trough, 300m) 
Escarpia spicata (Santa Catalina Basin, 1250m) 
ae Escarpia laminata (Florida Escarpment, 3300m) 
65 Escarpia spicata (Guaymas Seep, 1650m) 
Escarpia spicata (Guaymas Vent, 2000m) 
67 Riftia pachyptila Riftiidae 
Tevnia jerichonana 
Oasisia alvinae 
Ridgeia piscesae (Gorda Ridge) ER 
a Ridgeia piscesae (Juan de Fuca Ridge) é 
Ridgela piscesae (S Explorer Ridge) i 
Ridgeia piscesae (Escanaba Trough) : 
75 ,Lamellibrachia sp. (Nankai Trough,1200m) i 
100 (|, amellibrachia sp. (Nankai Trough, 2000m) ° 
Lamellibrachia columna (Lau Basin, 1850m) 
100 -Basibranchia sp. (Mariana Trough, 430m ) ? 
99 Lamellibrachia sp. (Nankai Trough, 300m) h 
93 Lamellibrachia barhami (Middle Valley, 2400m) i 
400 —Lamellibrachia barhami (Oregon Margin, 2000m) = 
pogonophoran sp. (Japan) 
Galathealinum brachiosum 
Nena alaaiagt 


0.02 


Fig. 6. Minimum evolution tree showing the relationships among tube worms based on a partial sequence 
(500 base pairs) of the mitochondrial cytochrome oxidase I gene, using the Kimura 2-parameter correction for 
multiple substitutions (references and Genbank accession numbers shown in Table 2). Numbers at nodes indicate 
the proportion of occurrences of a particular branching order in 1000 bootstrap replications. Bootstrap values 


below 65 not shown. GoM, Gulf of Mexico. 


mm beyond the anterior face of the obtu- 
raculum. While the medial structure of 
Seepiophila jonesi is prominent and readily 
visible, it does not project a great distance 
beyond the anterior face of the obturacu- 
lum. The greatest distance recorded in spec- 
imens available for this study was 3.5 mm. 
Also, the medial structure of S. jonesi ends 
bluntly, whereas those of the new genus 


(Southward et al. 2001) and E. spicata taper 
to a sharp point or terminate in a slightly 
bifid tip, respectively. 

Paired internal excretory ducts opening 
through a single pore is a feature Seepio- 
phila jonesi shares with other species in the 
Escarpiidae. However, in addition to having 
only a single type of filament associated 
with its branchial lamellae, the posteroven- 


704 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Collection locations, GenBank accession numbers and references for vestimentiferan specimens 


used in the present analysis of the COI gene. 


Organism Location GenBank accession Reference 
Escarpia laminata Florida Escarpment U74063 Black et al. 1997 
Escarpia sp. Nankai Trough (300 m) D50593 Kojima et al. 1997 
Escarpia sp. Nankai Trough (1200 m) D50594 Kojima et al. 1997 
Escarpia sp. Iheya Ridge (1400 m) D50595 Kojima et al. 1997 
Escarpia spicata Guaymas Basin seep U74065 Black et al. 1997 
Escarpia spicata Guaymas Basin vent U74064 Black et al. 1997 
Escarpia spicata Santa Catalina Basin whale fall U84262 Feldman et al. 1998 
Basibranchia sp. Mariana Trough U74078 Black et al. 1997 
Lamellibrachia barhami Middle Valley U74055 Black et al. 1997 
Lamellibrachia barhami Oregon margin U74054 Black et al. 1997 
Lamellibrachia columna Lau Basin U74061 Black et al. 1997 
Lamellibrachia sp. Nankai Trough (1200 m) D38029 Kojima et al. 1997 
Lamellibrachia sp. Nankai Trough (300 m) D38030 Kojima et al. 1997 
Lamellibrachia sp. Nankai Trough (2000 m) D50592 Kojima et al. 1997 
Oasisia alvinae 21°N Eastern Pacific Rise U74069 Black et al. 1997 
Seepiophila jonesi Gulf of Mexico, Bush Hill AF317287 This study 
Seepiophila jonesi Gulf of Mexico, GB425 AF317288 This study 
Seepiophila jonesi Gulf of Mexico, GC234 AF317289 This study 
Seepiophila jonesi Gulf of Mexico, TAMU-17 AF317290 This study 
Ridgeia piscesae Southern Explorer Ridge U74057 Black et al. 1997 
Ridgeia piscesae Juan de Fuca Ridge AF022233 Black unpublished 
Ridgeia piscesae Gorda Ridge U87978 Black et al. 1998 
Ridgeia piscesae Escanaba Trough AFO022235 Black unpublished 
Riftia pachyptila 9°N Eastern Pacific Rise U74074 Black et al. 1997 
Tevnia jerichonana 9°N Eastern Pacific Rise U74075 Black et al. 1997 
pogonophoran sp. Japan D50598 Kojima et al. 1997 
Galathealinum brachiosum Oregon AF178679 Boore & Brown 2000 
tral margin of the vestimentum is deeply Discussion 


incised in S. jonesi, whereas this margin is 
entire in species of Escarpia. Seepiophila 
jJonesi and species of Escarpia also differ 
in the relative position of the excretory 
pore. Jones (1985) states that the excretory 
pore is situated near the base of the obtu- 
raculum in Escarpia, whereas in S. jonesi 
it is positioned approximately one-third the 
distance from the base to the apex of the 
obturaculum. 

Collars associated with tubes is a wide- 
spread feature among basibranchian vesti- 
mentiferans, and therefore, we believe this 
characteristic should not be emphasized in 
comparative accounts. However, it is wor- 
thy of note that the tubes of Seepiophila 
jJonesi possess well-developed collars, 
whereas such collars are not present on 
tubes of species of Escarpia. 


Presently, 15 species of vestimentiferan 
tube worms have been recognized and de- 
scribed (Webb 1969; van der Land & 
Ngrrevang 1975; Jones 1981, 1985; Mané- 
Garzon & Montero 1985; Southward 1991; 
Miura et al. 1997; Southward & Galkin 
1997). Ridgeia phaeophiale, originally de- 
scribed by Jones (1985), was subsequently 
synonymized with R. piscesae by South- 
ward et al. (1995). Of the remaining 14 spe- 
cies, eight species are reported only from 
hydrothermal vent communities, including 
Riftia pachyptila, Oasisia alvinae, Tevnia 
jJerichonana, Ridgeia piscesae, Alaysia 
spiralis, Lamellibrachia columna, L. sat- 
suma and Arcovestia ivanovi. Four species 
are known exclusively from cold-water 
communities, including L. barhami, L. luy- 


VOLUME 114, NUMBER 3 


mesi, L. victori, and Escarpia laminata. Es- 
carpia spicata is reported from cold-water 
and hydrothermal vent sites (Jones 1985, 
Black et al. 1997). Seepiophila jonesi, at 
present, represents an additional species oc- 
curring only in cold-water communities. 

The recognition of Seepiophila jonesi 
raises the number of vestimentiferan spe- 
cies known from the Gulf of Mexico to 
three. Lamellibrachia cf. luymesi and S. 
jJonesi are sympatric in all of the shallow 
water hydrocarbon seep communities sam- 
pled for this study. The third species, Es- 
carpia laminata, is currently recorded only 
from deep water at the base of the Florida 
Escarpment (Jones 1985). 

In order to further clarify the phyloge- 
netic position of Seepiophila jonesi among 
other vestimentiferans, we conducted an 
analysis of the mitochondrial COI gene in 
specimens of the genera Seepiophila, Es- 
carpia, Riftia, Tevnia, Oasisia, Ridgeia, and 
Lamellibrachia (see Table 2). The average 
divergence for the COI gene among the 
four specimens of S. jonesi from the Loui- 
siana Slope was low (0.8%), whereas the 
average divergence among the vestimenti- 
ferans in the escarpid cluster was 7.6%. As 
a comparison, sequence divergence within 
the described group of L. barhami is 0.3%, 
and the divergence among the lamellibrach- 
ids is 4.5%. This difference in divergence 
is reflected in the deeper branches between 
species of the escarpid cluster than those of 
the lamellibrachid group (Fig. 6). The very 
low sequence divergence among the sam- 
ples of S. jonesi suggests that the four in- 
dividuals (found within 65 km of each oth- 
er) are members of a single, freely inter- 
breeding species. The very deep branching 
pattern between S. jonesi and other species 
in the family Escarpiidae is a reflection of 
the relatively high degree of sequence di- 
vergence between S. jonesi and the remain- 
ing species of the Escarpiidae. Although no 
definitive “‘molecular clock’’ exists that 
could generate benchmarks for the naming 
of genera, families, etc., the deep branching 
pattern within the Escarpiidae relative to 


705 


that within the Lamellibrachiidae supports 
the placement of the Louisiana Slope ves- 
timentiferans within a new genus but still 
within the larger cluster of the family Es- 
carpiidae. 


Acknowledgments 


This work was supported by the NOAA 
National Undersea Research Program at the 
University of North Carolina, Wilmington, 
Harbor Branch Oceanographic Institution, 
and the Minerals Management Service, 
Gulf of Mexico Regional OCS Office 
through the contract number 1435-10-96- 
CT30813. We thank the captain and crew 
of the RV Edwin Link as well as the sub- 
mersible crew and pilots of the Johnson Sea 
Link. We thank Erica Nix, Jim Childress, 
and Kim Nelson for collecting and provid- 
ing samples. We also thank W. R. Brown 
and S. Braden, Scanning Electron Micro- 
scope Laboratory, National Museum of 
Natural History, Smithsonian Institution, 
for the operation of their equipment. Dr. 
James Sidie and Dr. A. C. Allen kindly 
made SEM facilities available for our use 
at Ursinus College. Finally, we thank two 
anonymous reviewers for suggestions that 
greatly improved the manuscript. 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):708-719. 2001. 


Syllidae (Polychaeta) from San Quintin lagoon, Baja California, 


Mexico, with the description of a new genus 


Victoria Diaz-Castaneda and Guillermo San Martin 


(VD-C) Departamento de Ecologia, CICESE, Apartado Postal 2732, Ensenada, 
Baja California, Mexico; 


(GSM) Departamento de Biologia (Zoologia), Laboratorio de Biologia Marina e Invertebrados, 


Facultad de Ciencias, Universidad AutOnoma de Madrid, Espana 


Abstract.—During a recent study of the polychaetes from San Quintin lagoon 
(Baja California) 514 specimens belonging to the family Syllidae (Polychaeta) 
were collected, but only a few species were previously reported. In this paper, 
we report seven species: Syllis aciculata (Treadwell, 1945), Syllis gracilis 
(Grube, 1840), Syllis heterochaeta (Moore, 1909), Exogone (Exogone) lourei 
(Berkeley & Berkeley, 1938), Sphaerosyllis californiensis (Hartman, 1966), 
Grubeosyllis mediodentata, a new combination, and Cicese sphaerosylliformis, 
anew genus, and species. Grubeosyllis mediodentata is characterized by having 
cylindrical, somewhat elongated papilliform dorsal cirri, a long proventriculus 
and pharynx, pharyngeal tooth located in front of the midline of the pharynx, 
and compound setae with long bidentate blades provided with long, fine spines 
on margin. The new genus Cicese is similar to the genus Sphaerosyllis since 
it has antennae, tentacular cirri and anal cirri with bulbous bases, as well as 
papillae on the palps and dorsum, but it has two pairs of tentacular cirri instead 


of a single pair. 


The Syllidae is a very large family of 
polychaetes with a high number of de- 
scribed species; most species are small and 
usually overlooked in macrofaunal studies. 
Syllids of the Pacific coast of Mexico were 
reported in a few papers: Rioja (1941, 
1947a, 1947b, 1959, 1962); the papers of 
Rioja (1943), G6ngora-Garza (1984) and 
Gongora-Garza & de Leon Gonzdlez 
(1993) are the only ones devoted exclusive- 
ly to this family. The former paper also in- 
cludes a key for all the syllids known on 
the Pacific coast of México; a total of 49 
species of Syllidae is reported in the key. 
Other papers treating syllids from close ar- 
eas are those of Hartman (1968) and Ku- 
denov & Harris (1995) (for California), 
Hartmann-Schréder (1959) (for El Salva- 
dor), Fauchald (1977), Fauchald & Reiner 
(1975) and Lopez et al. (1997) (for Pana- 
ma) and Westheide (1974) (for the Gala- 


pagos Islands). Other studies dealing with 
syllids are Day (1967) and Fauchald & Rei- 
ner (1975). 

San Quintin lagoon is a highly produc- 
tive coastal lagoon located between 30°24’'— 
30°30'N and 115°57’—116°01'W on the Pa- 
cific coast of Baja California (Fig. 1). This 
lagoon has an area of 42 km? and around 
75% of it is covered by the eelgrass Zostera 
marina (Inclan-Rivadeneyra & Acosta-Ruiz 
1988, Ibarra-Obando 1990). It has been ex- 
ploited for many years (mariculture), but it 
can still be considered a relatively non-dis- 
turbed area, although intensive oyster mari- 
culture is being considered in the near fu- 
ture. 

The lagoon has the shape of an inverted 
“Y”’?, the western arm (Falsa Bay) has an 
average depth of 4 m, whereas the eastern 
arm (San Quintin Bay) has an average 
depth of 8 m. Granulometric studies show 


VOLUME 114, NUMBER 3 


Kenton Hill 


Mt. Ceniza 


: Mt Mazo 


0 s00 1000 
METROS 


2000 
116°00' 


Fig. 1. 


that in shallow areas, as well as to the north 
of both arms, clay and silty-sand predomi- 
nate, whereas near the mouth very fine 
sands are more abundant. The channel sed- 
iments are highly diverse, ranging from me- 


709 


e 
Bahla 


de @22 


: San Quintin 
° e 


oy je 
20 
23, 


115°S5'"W 


Study site: sampling stations in San Quintin Bay, Baja California, Mexico. 


dium to fine sand and silt (Barnard 1970, 
Calderon-Aguilera 1992). The lagoon mar- 
gins present a typical saltmarsh flora with 
Spartina foliosa and Salicornia virginica 
among other vascular plants. Syllids are 


710 


more abundant in areas of fine sand, cov- 
ered by Zostera marina and Spartina folio- 
Sa. 


Materials and Methods 


Forty-four stations distributed in both 
arms were sampled in December 1995 and 
April 1998 (Table 1). Two replicates per 
station were collected with a geological 
corer (16 cm internal diameter, 12 cm 
depth) with a sampling area of 0.02 m/. 
Sediments were sieved in the field using 1 
mm mesh size and retained material was 
preserved in 7% buffered formaldehyde. In 
the laboratory, samples were washed using 
a 0.5 mm mesh and transferred to 70% iso- 
propanol. Polychaetes were then sorted and 
syllids identified to species level. 

In the material examined, BSQ represent 
the initials for Bahia San Quintin, followed 
by the station number. Samples are repre- 
sented by M and replicates by R. Speci- 
mens are deposited in the Polychaete Col- 
lection of Universidad Aut6noma de Nuevo 
Leé6n (UANL). 


Results 


Family Syllidae Grube, 1850 
Subfamily Syllinae Grube, 1850 
Genus Syllis Savigny in Lamarck, 1818 
Syllis aciculata Treadwell, 1945 


Typosyllis aciculata Treadwell, 1945:1—2, 
figs. 1-5; Hartman, 1968:475, text-figs. 
1-7. 


Material examined.—BSQ 42R (6). 
Distribution.—From California to Pana- 
ma; South Chinese Sea. 


Syllis gracilis Grube, 1840 


Syllis gracilis Grube, 1840:77, pl. 31a; 
San Martin, 1984:376, pls. 97, 98. 

Syllis (Syllis) gracilis—Fauvel, 1923:259, 
fig. 96F—I; Day, 1967:241, fig. 12.1m—p. 


Material examined.—BSQ 14 (1), BSQ 
7M (1), BSQ 7R (1). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Distribution.—Cosmopolitan in temper- 
ature and tropical seas. 


Syllis heterochaeta Moore, 1909 


Syllis (Ehlersia) heterochaeta.—Rioja, 
1941:694—-695; Hernandez-Alcdntara, 
1992:200. 

Langerhansia heterochaeta.—Hartman, 
1968:434—435, figs. 1-7. 


Material examined.—BSQ 43M (5). 
Distribution.—Western Mexico to west- 
ern Canada. 


Subfamily Exogoninae Rioja, 1925 
Genus Grubeosyllis Verrill, 1900 
Grubeosyllis mediodentata (Westheide, 
1974), new combination 
Figs. 2—4 


Brania mediodentata Westheide, 1974:93— 
97, figs. 42A—C, 43; Russell, 1991:52— 
54, Fig. 2. 


Material examined.—BSQ 29 (1). BSQ 
6R (1). BSQ 19R (2). BSQ 23 (1). BSQ 39 
(3). BSQ R15 (1). BSQ R16 (1). BSQ 27R 
(1). BSQ 17R (1). BSQ 6 (1). BSQ 28R 
(1). BSQ 25 (1). BSQ 19 (2). BSQ 14 (18) 
+ 2 specimens used for SEM observations. 

Description.—Body small, slender, 1.9 
mm long, 0.2 mm wide, for 31 setigers, 
without color markings; intersegmental fur- 
rows well marked (Fig. 2A). Prostomium 
ovate, about 2.2 times wider than long; two 
pairs of lensed eyes in open trapezial ar- 
rangement and two small anterior eyespots. 
Antennae elongate, fusiform, with small 
subdistal enlargement, shorter than prosto- 
mium and palps together; median antenna 
of same length or slightly longer than pro- 
stomium, originating in middle of prosto- 
mium; lateral antennae somewhat shorter 
than median antenna, originating in front of 
anterior eyes, slightly posterior to eyespots 
(Figs. 2A, B, 3A). Palps small, shorter than 
prostomium, fused along entire length, with 
distal notch. Tentacular segment well de- 
fined, similar in length to following seg- 
ments; two ciliated lateral nuchal organs 


VOLUME 114, NUMBER 3 


Table 1.—Station locations and granulometry at San Quintin lagoon, Baja California. 


Station Lat. N. Long. W. Eh(mV) 
1 30°29'20" 115°59'35” 
2 30°29'34” 115°59’34" = 255) 
3 30°29'06" 115°59’02” 
4° 30°29'23” 115°58'57" —184 
5 30°29'06" 115°59'01” SNZy 
6 30°29'02” 115°58’37" 103 
7 30°28'38" 115°58'26" —150 
8 30°28'45” 115°58’10" —280 
9 30°28'50” Liss7 si” = 37) 
10 30°28'28" 115°57'29” 168 
11 30°28'18” 115°57'46" —294 
12 30°27'52” 115°57'50" —145 
13 30°27'48" 115°57'46” —203 
14 30°27'49” 115°57'30" —320 
15 30°28'02” 115°57’11" —126 
16 30°27'33" 115°57'41" —360 
17 30°27'21” 115°57/09" —230 
18 30°27'46" 115°57'06" —200 
19 30°26'32” ISS? 0” S252 
20 30°26'46" 115°57’00" —156 
21 30°26'52” 115°56’37" = 27] 
a) 30°27'07" 115°56/17" —30 
23 30°26'33” 115°56'03” 152 
24 30°25'52” MIS sy” 
5) 30°26'22” 115°56’33" =25)]/ 
26 30°25'58” 115°56’48” —166 
27 30°25'50" 115°57’09" = 783} 
28 30°25'12” 115°57'31” —136 
29 30°24'32” 115°57'29" —242 
30 30°24'29" 115°58'38" —94 
31 30°25'10” 115°58'12” —104 
32 30°24'39” 115°59'46" —174 
33 30°25'12” 115°59'44" —285 
34 30°25'32” 115°00'14" —84 
35 30°25'42” 115°59'44" —326 
36 30°25'52” 115°59'12” —104 
37 30°26'21” 115°59'33” —205 
38 30°26'10" 115°59’58" —115 
39 30°26'00" 115°00'26” —148 
40 30°26'32” 115°00'36" 181 
4] 30°26'42” 115°00'42” 100 
42 30°26'54” 115°59'55” —86 


between prostomium and peristomium (Fig. 
3A); two pairs of tentacular cirri, short, 
smaller than antennae, ovate to conical, 
somewhat larger at bases (Figs. 2A, B, 3A, 
B). Dorsal cirri on all setigers; anterior dor- 
sal cirri similar in shape to dorsal tentacular 
cirri but somewhat longer, shorter than 
parapodial lobes, but progressively longer 
and slender with very small difference in 


711 

TES O.M. (%) Sand (%) Silt (%) Clay (%) 

De3X5) 39.86 30.68 29.28 
21.4 0.43 96.8 3.2 0) 

1.86 63.78 23.64 7.9 
20.8 1.33 60.08 25.81 13.61 
20.1 2.81 44.97 30.12 22.46 
20.5 2.78 13.42 65.52 21.26 
20.5 1.91 63.79 23.38 10.21 
20.4 0.58 71.88 27.84 0.77 
20.6 2.66 36.2 36.38 22.78 
21.4 Dred I] 65.82 18.78 14.63 
20.5 0.79 84.38 11.51 4.11 
19.1 0) 25.84 66.09 8.1 
20.5 0.58 86.72 13.28 0 
21 1.55 50.59 35.64 12.8 
19.9 333) 19.77 44.88 27.69 
20 3.1 96.68 3.32 0) 
20.4 1.89 34.51 31.55 14.86 
20.4 1.4 69.61 6.59 ANS 
21.4 2.03 60.08 25.81 13.61 
20.1 0.62 74.65 25.04 2.36 
20.3 0.76 78.61 12.34 9.26 
18.9 1.68 37.77 35.52 22.97 
19.3 1.27 42.58 40.04 25.89 

2.23 31.95 28.86 24.06 
20.7 0.89 52.38 24.01 20.78 
21.6 0.43 97.06 DAS 0 
21.1 1.38 98.08 1.92 0) 
20.5 0.48 89.05 10.95 0 
21.6 0.28 98.13 Deda) 0 
20.5 0.05 71.78 17.04 8.86 
21.5 0.37 88.4 11.79 0 
20.5 1.99 66.82 21.8 10.39 
20.6 0.26 93.75 6.25 0 
21.1 2.96 51.3 36.54 12.52 
21.1 2.21 42.27 45.83 6.4 
21.5 0.53 58.99 33.54 7.03 
20.6 0.93 30.03 60 6.05 
21 Del 39 46.14 IS27/ 
19.8 1.16 7.26 72.59 8.17 
21.9 
21.6 
21.3 


width between bases and tip, cylindrical 
(Fig. 2A, B, D); dorsal cirri of midbody 
somewhat longer than parapodial lobes 
(Fig. 2C); from midbody posteriorly, dorsal 
cirri progressively smaller. Parapodial lobes 
conical, with presetal papilla (Fig. 2C). 
Ventral cirri digitiform, shorter than para- 
podial lobes. Compound setae hetero- 
gomph, with short, fine subdistal spines on 


WZ 


shafts, and blades strongly bidentate, both 
teeth similar in length and shape or proxi- 
mal tooth somewhat shorter than distal 
tooth, well separated from each other, elon- 
gate, provided with long, upwardly project- 
ing spines on margin, especially on most 
anterior and dorsal setae (Figs. 2E, H, 4A— 
C). Anterior parapodia each with about 10 
compound setae, with marked dorsoventral 
gradation, 28 ym blades on dorsalmost se- 
tae, 18 ym blades on ventralmost ones (Fig. 
2E); progressively decreasing number of 
compound setae on each parapodium, pos- 
terior parapodia each with about 5 com- 
pound setae, blades 32 wm dorsally, 14 wm 
ventrally (Fig. 2H). Solitary dorsal simple 
seta from anterior parapodia, thick, strongly 
bidentate, with large, triangular proximal 
tooth, forming about a 45° angle with the 
distal tooth, smooth on anterior parapodia 
(Fig. 2F), progressively provided with more 
spines on margin (Fig. 21), forming several 
rows of spines (observations by SEM) (Fig. 
4D). Solitary ventral simple seta on each 
posterior parapodium, sigmoid, similar to 
dorsal simple seta but slender, smooth (Fig. 
2J) or provided with few short spines on 
margin (SEM observations) (Fig. 4E). An- 
terior parapodia each with two slender acic- 
ulae, one straight and other acuminate (Fig. 
2G), provided with subdistal enlargement 
and conical tip; single aciculum on each 
middle and posterior parapodium, acumi- 
nate (Fig. 2K). Pharynx long and wide, 
through about 5—6 segments (Fig. 2A); 
mouth opening provided with circle of cilia 
and about 10 soft, very small papillae (Fig. 
3B); pharyngeal tooth very small, incon- 
spicuous, conical, located just in front of 
middle of pharynx, far from anterior margin 
of pharynx (Fig. 2A, B). Proventriculus 
similar in length to pharynx, through about 
5 segments, with about 27 rows of muscle 
cells (Fig. 2A). Females carrying eggs on 
dorsolateral position of midbody; eggs at- 
tached to body by means of thin notosetae 
Gis. S© 3D): 

Remarks.—Most of the species of the ge- 
nus Grubeosyllis have fusiform dorsal cirri, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


with bulbous bases and long, pointed tips; 
however, a few species of this genus have 
cylindrical, digitiform dorsal cirri, as in G. 
mediodentata. This species has a body very 
similar to G. swedmarki (Gidholm, 1962), 
from the North Atlantic, but the compound 
setae are completely different being short 
and unidentate in G. swedmarki (Gidholm 
1962, Parapar et al. 1993) and long and bi- 
dentate in G. mediodentata. Grubeosyllis 
celiae (Parapar & San Martin, 1992), from 
Ceuta (North West Africa) has longer dorsal 
cirri, a longer proventriculus, compound se- 
tae with shorter blades than those of G. me- 
diodentata and the dorsal simple setae are 
unidentate (Parapar & San Martin 1992). 

This species reproduces by external ges- 
tation, as many others of the same genus 
and the subfamily Exogoninae. The eggs 
are attached to the body by means of epi- 
tokous setae (Fig. 3C), as recently reported 
by Kuper & Westheide (1998) for other 
similar species. 

Distribution.—Galapagos Islands, Be- 
lize. This species is newly recorded for 
Mexico. 


Cicese, new genus 


Diagnosis.—Body small, cylindrical. 
Prostomium with two pairs of lensed eyes 
and two eyespots. Palps fused along entire 
length. Peristomium well defined, not fused 
to prostomium, with two pairs of tentacular 
cirri. Dorsal and ventral cirri on all para- 
podia. Antennae, tentacular cirri, anal cirri 
and, at least, some dorsal cirri with bulbous 
bases and slender tips. Palps, pygidium and 
dorsum provided with papillae. Two anal 
cirri. Pharynx provided with an anterior 
middorsal tooth and a crown of papillae. 
Parapodia each with several compound se- 
tae, and dorsal and ventral simple setae on 
some parapodia. Aciculae acuminate. Fe- 
males carrying eggs dorsally by means of 
notosetae and mature males provided with 
long natatory setae. 

Remarks.—The new genus is close to a 
group of species of the genus Sphaerosyllis, 


VOLUME 114, NUMBER 3 713 


Fig. 2. Grubeosyllis mediodentata (Westheide, 1974). A, anterior end, dorsal view. B, detail of prostomium 
and anterior setigers. C, midbody parapodium. D, posterior end, dorsal view. E, compound setae, anterior para- 
podium. E dorsal simple seta, anterior parapodium. G, aciculae, anterior parapodium. H, compound setae, pos- 
terior parapodium. I, dorsal simple seta, posteriro parapodium. J, ventral simple seta. K, aciculum, posterior 
parapodium. Scale: A, D—0.14 mm; B—0.6 mm; E~-K—15 pm. 


714 


Magn ee. Fi 


50x bua . 


Fig. 3. 


having more or less elongate dorsal cirri on 
the midbody, two pairs of eyes and a pair 
of eyespots on the prostomium, acuminate 
aciculae, compound setae with bidentate 
blades and small, inconspicuous papillae on 
the dorsum, palps and pygidium (e.g., 
Sphaerosyllis bilobata Perkins, 1981, S. 
cryptica Ben-Eliahu, 1977, S. belizensis 
Russell, 1989, and others) (see Perkins 
1981, Ben-Eliahu 1977, Russell 1989). 
However, all these species, as members of 
the genus Sphaerosyllis, are provided with 
a single pair of tentacular cirri. On the other 
hand, Cicese is also closely related to Gru- 
beosyllis (Verrill, 1900), sharing the shape 
of the aciculae, similar setae, and having 
two pairs of tentacular cirri. However, Ci- 
cese has papillae, whereas Grubeosyllis has 
a smooth dorsum, palps and pygidium, a 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ig al : 
iz, : vie 
OUUN -. a 


Grubeosyllis mediodentata (Westheide, 1974). SEM. A, detail of prostomium, dorsal view. B, later- 
oventral view with proboscis everted. C, an egg, attached to body by means of capillary notosetae. D, detail of 
capillary notosetae. 


character considered as exclusive to Sphae- 
rosyllis. 

Etymology.—The name is given in honor 
of the research center CICESE (Centro de 
Investigaci6n Cientifica y Estudios Super- 
iores de Ensenada) located in Baja Califor- 
nia, Mexico. 


Cicese sphaerosylliformis, new species 
Fig. 5 


Material examined.—BSQ R§8; Holo- 
type. BSQ R17; 1| paratype. BSQ 23; | par- 
atype. 

Description.—Body small, short, holo- 
type complete mature female 1.5 mm long, 
0.2 mm wide, for 29 setigers, without color 
markings; dorsum covered with very small 
papillae, longer on posterior part of body 


VOLUME 114, NUMBER 3 


715 


Fig. 4. Grubeosyllis mediodentata (Westheide, 1977). SEM. A, dorsal compound seta. B, median compound 
seta. C, ventral compound seta. D, dorsal simple seta. E, ventral simple seta. 


(Fig. 5D, E). Prostomium ovate, more than 
twice as wide as long (Fig. 5A); two pairs 
of large, lensed eyes in very open trapezoi- 
dal arrangement, nearly on line, anterior 
pair larger than posterior pair, and two an- 
terior eyespots. Median antenna longer than 


prostomium and palps together, large at 
base, enlarged at midlength and with slen- 
der tip, originating between posterior eyes; 
lateral antennae onion-shaped, with bulbous 
bases and short, slender tips, shorter than 
median antenna, originating close to eye- 


716 


spots. Palps short, fused along entire length, 
with indistinct dorsal furrow, provided with 
few very small, papillae dorsally. Tentacu- 
lar segment shorter than following seg- 
ments, well defined, covering dorsal poste- 
rior part of prostomium, dorsally bilobed; 
dorsal tentacular cirri similar in length to 
median antenna, similar in shape to lateral 
antennae but somewhat more elongate (Fig. 
5A); ventral tentacular cirri similar in shape 
to lateral antennae, but somewhat smaller 
(Fig. 5B). Dorsal cirri of setiger one similar 
in shape to dorsal tentacular cirri, somewhat 
longer. Dorsal cirri of setigers 2 and 3 
small, with strongly bulbous bases and 
short, slender tips. Dorsal cirri from setiger 
3 elongate, with slightly enlarged bases and 
long, slender tips (Fig. 5C), progressively 
longer and slender; dorsal cirri on midbody 
only somewhat shorter than body width 
(Fig. SA); dorsal cirri from mid-posterior 
setigers shorter, less elongate, with wider 
bases (Fig. 5D). Parapodial lobes short, 
conical; ventral cirri digitiform, shorter than 
parapodial lobes (Fig. 5C). Compound setae 
heterogomph, provided with slender, biden- 
tate blades, proximal tooth smaller than dis- 
tal tooth, with long, thin, erect basal spines 
on margin, progressively shorter, smooth 
distally (Fig. 5H, I). Anterior parapodia 
each with about 10—13 compound setae, 
blades 34 wm dorsally, 16 wm ventrally; 
progressively reduced number of compound 
setae, on each parapodium to 6—7 on pos- 
terior parapodia, similar to anterior com- 
pound setae. Solitary dorsal simple seta 
from anterior parapodia (from setiger 3 in 
holotype), slender, unidentate, provided 
with short, thin spines on margin (Fig. 5G). 
Solitary ventral simple seta on far posterior 
parapodia, slender, sigmoid, bidentate, 
smooth. Acicula solitary, slender, acumi- 
nate, provided with a subdistal enlargement 
and a long, filiform tip (Fig. 5F). Pharynx 
slender, everted in holotype (Fig. 5A), with 
pharyngeal tooth located anteriorly, sur- 
rounded by crown of about 10 small soft 
papillae; a few, probably 5, subdistal papil- 
lae on pharynx (Fig. 5B). Proventriculus 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


shorter than pharynx, through about 4 seg- 
ments, with about 17 roes of muscle cells. 
Pygidium small, provided with small papil- 
lae and two long anal cirri, with bulbous 
bases, longer than posterior dorsal cirri 
(Fig. 5D). Holotype carrying eggs dorsally 
from setiger 11; one paratype mature male, 
provided with natatory setae from setiger 
10. 

Remarks.—Rioja (1943) described Bran- 
ia limbata arenacea from the Pacific coast 
of México with two pairs of tentacular cirri 
and the dorsum covered by detritus, a char- 
acter of the genus Sphaerosyllis. Apparent- 
ly, Brania limbata arenacea (Rioja, 1943) 
is a member of Cicese, new genus. How- 
ever, it differs from Cicese sphaerosyllifor- 
mis, new species, by having antennae, ten- 
tacular cirri and dorsal cirri all similar, with 
small differences in lengths, unidentate 
blades of compound setae and lacking eye- 
spots. 

Etymology.—Meaning the form of this 
species resembles that of Sphaerosyllis. 


Genus Sphaerosyllis Claparéde, 1863 
Sphaerosyllis californiensis Hartman, 1966 


Sphaerosyllis_ californiensis.—Kudenov & 
Harris, 1995:28—30, Fig. 1.8. 


Material examined.—BSQ 6M (1). BSQ 
7R (1). BSQ 8R (1). BSQ 11M (1). BSQ 
12M (1). BSQ 14M (4). BSQ 14R (1). 21M 
(2). BSQ 26R (1). BSQ 28M (2). BSQ 31M 
(1). 

Distribution.—Southern California to 
Baja California. 


Genus Exogone Orsted, 1845 
Subgenus Exogone Orsted, 1845 
Exogone (Exogone) lourei Berkeley & 
Berkeley, 1938 


Exogone (Exoegone) lourei San Martin, 
1991:735—737; Kudenov & Harris, 1995: 
15/5 bicasleo. 


Material examined.—BSQ 3R (6). BSQ 


5M (9). BSQ 5R (5). BSQ 6M (47). BSQ 
6R (4). BSQ 8M (2). BSQ 8R (14). BSQ 


VOLUME 114, NUMBER 3 717 


Fig. 5. Cicese sphaerosylliformis, new genus, new species. A, anterior end, dorsal view. B, anterior end, 
ventral view. C, mid-posterior parapodium. D, posterior end, dorsal view. E, detail of surface of dorsum, mid- 
posterior segments. E aciculum. G, dorsal simple seta. H, compound setae, anterior parapodium. I, compound 
setae, posterior parapodium. A, B, F—I, from holotype; C—E, from paratype. Scale: A-E—0.65 mm; F—I—15 
pm. 


718 


9M (2). BSQ 9R (2). BSQ 11R (4). BSQ 
12M (1). BSQ 12R (48). BSQ 13R (4). 
BSQ 14M (16). BSQ 14R (17). BSQ 15R 
(18). BSQ 16M (8). BSQ 16R (16). BSQ 
17M (56). BSQ 17R (104). BSQ 18R (1). 
BSQ 19R (4). BSQ 20M (1). BSQ 20R (4). 
BSQ 21M (1). BSQ 21R (2). BSQ 22M (1). 
BSQ 22R (12). BSQ 23M (1). BSQ 23R 
(3). BSQ 24R (1). BSQ 25M (1). BSQ 25R 
(7). BSQ 26M (2). BSQ 26R (5). BSQ 27M 
(1). BSQ 27R (2). BSQ 34M (4). BSQ 37R 
(6). BSQ 39M (6). BSQ 39R (6). 

Distribution.—Northeast Pacific, from 
British Columbia to México. Gulf of Méx- 
ico, Cuba, Florida and Canary and Cape 
Verde Islands. 


Acknowledgments 


We would like to thank M. Necocechea 
and G. de la Selva for helping sort the ma- 
terial. Financial and technical support was 
provided by Centro de Investigaci6n Cien- 
tifica y Estudios Superiores de Ensenada 
(Baja California, Mexico). 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):720—724. 2001. 


Recognition of Cenogenus Chamberlin, 1919 (Polychaeta: 
Lumbrineridae) based on type material 


Luis F Carrera-Parra 


El Colegio de la Frontera Sur, Unidad Chetumal. Adpo. Postal 424, 
Chetumal QR. 77000, Mexico 


Abstract.—Cenogenus Chamberlin with C. descendens Chamberlin as its 
only species has been considered a junior synonym of Lumbrineris. It is here 
redescribed and emended as a distinct genus. Its diagnostic features include a 
single antenna in the nuchal fold; single digitate branchiae on anterior para- 
podia; maxillary apparatus of labidognath type, with four plates, maxillae II 
and IV edentate, maxillae V absent; mandible partially fused and simple mul- 
tidentate hooded hooks. Paraninoe Levenstein (type species Ninoe fusca 
Moore) is a junior synonym of Cenogenus; both species, C. descendens and 
C. fusca are recognized based on type material. 


Previously the external characters in lum- 
brinerids were considered to be uniform 
with simplicity and reduction of many mor- 
phological parts. Consequently, we have 
had a simplified generic system for the fam- 
ily, lumping all species described into only 
three or four genera. Current work on the 
taxonomy of lumbrinerids is changing this 
view; however, as a consequence of our ear- 
lier conceptions of the group, some genera 
were regarded as junior synonyms and have 
been forgotten or not considered during the 
creation of new taxa. 

Chamberlin (1919) erected the genus 
Cenogenus to include specimens collected 
in abyssal depths provided with ‘“‘a conical 
nuchal process present at anterior edge of 
first segment above, with four pairs of max- 
illae,”’ separating it in his key from other 
genera by the presence of only simple 
hooded hooks and with maxillae III and IV 
edentate. Hartman (1944), based upon these 
features, considered it a valid genus. How- 
ever, Fauchald (1970) regarded this genus 
as a junior synonym of Lumbrineris de 
Blainville. Levenstein (1977) erected Par- 
aninoe to include species provided with a 
nuchal organ and simple digitate branchiae; 


she noted that most of the included species 
were confined to abyssal depths. 

Here, I redescribe and emend Cenogenus 
and regard Paraninoe Levenstein, 1977, as 
a junior synonym. 


Materials and Methods 


Type materials were borrowed from the 
collections of the National Museum of Nat- 
ural History (USNM), Smithsonian Insti- 
tution, Washington, and the Museum of 
Comparative Zoology (MCZ), Harvard 
University, Cambridge. The measurements 
were standardized to setiger 10; they are 
abridged as L10 for length to setiger 10, 
and W10 for width at setiger 10. Ilustra- 
tions were made with a camera lucida. 


Cenogenus Chamberlin, 1919, emended 


Cenogenus Chamberlin, 1919:333—334. 
Paraninoe Levenstein, 1977:189—197, figs. 
ES 


Type species.—Cenogenus descendens 
Chamberlin, 1919, by original designation. 
Emended diagnosis.—Single small an- 
tenna in nuchal fold. Setae include limbate 
capillaries, limbate robust, and simple mul- 


VOLUME 114, NUMBER 3 


tidentate hooded hooks. Anterior segments 
with a parapodial branchia dorsal and pos- 
terior to parapodia. Maxillary apparatus and 
labidognath type; with four pairs of maxil- 
lae, maxillae I forceps-like with smooth 
edges and bridles poorly developed, max- 
illae II of similar length to maxillae I, max- 
illae III and IV edentate plates (maxillae V 
absent); mandibles partially fused. 


Discussion 


Chamberlin (1919) erected Cenogenus 
based on the presence of a nuchal antenna, 
maxillary apparatus with four well devel- 
oped plates with maxillae III and IV eden- 
tate and setae limbate and simple multiden- 
tate hooded hooks. 

Chamberlin misinterpreted the parapodial 
shape of C. descendens; he regarded the 
parapodia as lacking a presetal lobe and 
having a finger-like postsetal lobe. In fact, 
the parapodia have both lobes, but they are 
inconspicuous, and there is a simple digitate 
branchia posterior to the parapodia on the 
dorsal side in anterior segments. 

Fauchald (1970), following the tradition- 
al classification, regarded these characters 
as present in some species of Lumbrineris, 
and thus considered Cenogenus as a junior 
synonym of Lumbrineris. However, Lum- 
brineris, as defined in older systematic 
works, is a heterogeneous taxon that would 
include any abranchiate lumbrinerid; Lum- 
brineris s.s. includes only species without 
nuchal antennae and branchiae, with five 
pairs of maxillae and both simple and com- 
posite multidentate hooded hooks present 
(Orensanz 1990). Because the maxillary ap- 
paratus has four plates, anterior parapodia 
have a single branchia, nuchal antenna is 
present and only simple multidentate hood- 
ed hooks occur, Fauchald’s synonym is here 
considered erroneous and Cenogenus is a 
distinct valid genus. 

Levenstein (1977) erected Paraninoe to 
include species with a nuchal antenna, max- 
illary apparatus with four plates and postse- 
tal lobe with a simple digitate branchia; 


721 


most were formerly included in Ninoe Kin- 
berg. She stated that Paraninoe differs from 
Ninoe in the number of branchial filaments 
and in the shape of maxillae III and IV and 
listed five species included in her new ge- 
nus. Later, Orensanz (1990) increased this 
list to 10 species. 

Examination of type material of Ceno- 
genus descendens Chamberlin, 1919 and 
Ninoe fusca Moore, 1911 (type species of 
Paraninoe) revealed that Cenogenus is a 
valid genus and Paraninoe Levenstein, 
1977 is a junior synonym of it. 

Following the list of species provided by 
Levenstein (1977) and Orensanz (1990), the 
species of Cenogenus includes C. abyssalis 
(Imajima & Higuchi, 1975), C. antarctica 
(Monro, 1930), C. brevipes (McIntosh, 
1903), C. descendens Chamberlin, 1919, C. 
fusca (Moore 1911), C. fuscoides (Fau- 
chald, 1970), C. hartmanae (Levenstein, 
1977), C. monotentaculata (Averincev, 
1972), C. nagae (Gallardo, 1968), and C. 
simpla (Moore, 1905). 

Lumbrinereis minuta Théel, 1879 was 
placed as a member of Paraninoe by Miura 
(1980); in the original description, its max- 
ilary apparatus was described without 
maxillae V and only simple hooded hooks. 
However, Oug (1998) reviewed the type 
material and stated that it has a maxillary 
apparatus with five pairs of maxillae, with 
maxillae V free. Oug also indicated the 
presence of more than one species in the 
type material of Théel’s species. A study of 
better material is needed to assess its ge- 
neric status. 

Consequently, Lumbrinereis minuta 
Théel, 1879 cannot belong to Cenogenus; 
however, P. minuta sensu Miura (1980) is 
a species of Cenogenus. 


Cenogenus descendens Chamberlin, 1919 
Fig. 1A-—G 


Cenogenus descendens Chamberlin, 1919: 
333-334. 


Material examined.—Syntypes of Ceno- 
genus descendens Chamberlin (USNM 


722 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


G 
| C E 


Fig. 1. Cenogenus descendens Chamberlin, 1919. A, prostomium, dorsal view (USNM 19344); B, second 
parapodium, frontal view (USNM 19344); C, long limbate seta (MCZ 2302); D, stout limbate setae (MCZ 2302); 
E, simple multidentate hooded hook (MCZ 2302); EK maxillary apparatus (USNM 19344); G, mandibles (USNM 
19344); Cenogenus fusca (Moore, 1911); H, second parapodium in frontal view (USNM 17338). Scales: A = 
15 mm; B, C, G, H = 0.2 mm; D-E = 13 pm; F = 26 pm. 


19344, one specimen and MCZ 2302, one Description.—Syntypes of C. descen- 
specimen and one slide). Peru, 111 miles dens are all incomplete. USNM specimen 
NW off Aguja Point (5°42’S, 83°0'’W), 11 broken in four fragments, anteiror end with 
Nov 1904, Albatross sta. 4651, 4066 m. 29 setigers (LIO = 5.2 mm, W10 = 2.5 


VOLUME 114, NUMBER 3 


mm); MCZ specimen broken in two parts 
with ca. 70 setigers (LIO = 5.1 mm and 
W10 = 2 mm). 

Prostomium conical, short, about as long 
as wide, with a pair of divergent dorsal lon- 
gitudinal black bands; with a pair of nuchal 
organs and an antenna in the nuchal fold 
(Fig. 1A). Peristomium shorter than prosto- 
mium; separation between rings distinct 
dorsally and laterally, ventrally with a shal- 
low lip; both rings of similar length. 

All parapodia well developed, but first 
four smaller. Parapodia with inconspicuous 
pre- and postsetal lobes. First parapodia 
with an unilobulated digitate branchia at- 
tached dorsal and posterior to the parapodia 
(Fig. 1B), diminishing in length on middle 
segments and absent on the posterior seg- 
ments. 

Anterior parapodia with very long lim- 
bate setae (Fig. 1C), middle and posterior 
parapodia with stout limbate setae, shorter 
than setae of anterior parapodia (Fig. 1D), 
and simple multidentate hooded hooks (Fig. 
1E). All setae with black core from the base 
to near tip where they become translucent. 
Parapodia with three to five black aciculae 
with mucro; aciculae located in the middle 
of parapodia in all setigers; straight. 

Short and stout maxillary apparatus, with 
four pairs of maxillae (Fig. 1F). Mandibles 
well calcified, with shaft separated along 
half of length (Fig. 1G). Maxillary carriers 
distinctly shorter than maxillae I, anterior 
end slightly constricted. Maxillae I forceps- 
like with smooth edges, bridles poorly de- 
veloped; maxillae II stout, of similar length 
to maxillae I, with two broad teeth in the 
anterior end; maxillae III and IV edentate 
plates. 

Distribution.—Off Aguja Point, Peru, in 
abyssal depth. 


Cenogenus fusca (Moore, 1911) 
Fig. 1H 


Ninoe fusca Moore, 1911:285—288, Pl. 19, 
Figs. 110-118. 
Paraninoe fusca Levenstein 1977:190—-191. 


723 


Material examined.—Holotype of Ni- 
noe fusca, Moore (USNM_ 17338), off 
Santa Catalina Islands, California, U.S.A. 
(33°10'15"N, 121°42’15"W) 1 Apr 1904, 
Albatross Sta. 4397, 3953 m. 

Description.—Specimen broken in two 
fragments, anterior end with 45 setigers 
(L10 = 6.3 mm and W10 = 3.2 mm). 

The holotype of Ninoe fusca lacks max- 
illary apparatus, but it was well described 
and illustrated by Moore (1911) with four 
pairs of maxillary plates, maxillae II biden- 
tate and maxillae III and IV edentate. 

The shape of prostomium, peristomium, 
parapodia, setae and the distribution of the 
branchiae and setae resemble those of C. 
descendens. However, in parapodia two to 
five the aciculae are located in dorsal po- 
sition and are curved (Fig. 1H). 

Distribution.—Off Santa Catalina Is- 
lands, California, U.S.A. in abyssal depth. 


Acknowledgments 


I wish to thank Ardis B. Johnston (Mu- 
seum of Comparative Zoology, Harvard 
University) and Kristian Fauchald (National 
Museum of Natural History, Washington, 
D.C.) for making available the type mate- 
rials that made this study possible, and for 
making laboratory facilities available. I am 
grateful to Leonard P. Hirsch and K. Fau- 
chald for housing me during my visit to the 
NMNH. I also wish to thank Sergio I. Sa- 
lazar-Vallejo, Kristian Fauchald and José 
M. Orensanz for discussions and help in 
preparation of the final version of the man- 
uscript. This study was partially financed by 
a research fellowship from El Colegio de la 
Frontera Sur and by CONACYyT (32529-T). 


Literature Cited 


Averincev, V. G. 1972. Benthic polychaetes Errantia 
from the Antarctic and Subantarctic collected 
by the Soviet Antarctic Expedition._Akademia 
Nauk SSSR, Zoologisheskii Institut Issledocan- 
ia Fauna Morei 11:88—293 (in Russian). 

Chamberlin, R. V. 1919. The Annelida Polychaeta.— 
Memoirs of the Museum of Comparative Zo- 
ology at Harvard College 48:1—514. 


724 


Fauchald, K. 1970. Polychaetous annelids of the fam- 
ilies Eunicidae, Lumbrineridae, [phitimidae, Ar- 
abellidae, Lyrsaretidae and Dorvilleidae from 
Western Mexico.—Allan Hancock Foundation 
Monographs in Marine Biology 5:1—335. 

Gallardo, V. A. 1968. Polychaeta from the Bay of Nha 
Trang, South Viet Nam.—Naga Report, Scien- 
tific Results of Marine Investigations of the 
South China Sea and Gulf of Thailand 1959— 
1961 4:35—279. 

Hartman, O. 1944. Polychaetous annelids, 5. Euni- 
cea.—Allan Hancock Pacific Expeditions 10:1— 
238. 

Imajima, M., & M. Higuchi. 1975. Lumbrineridae of 
polychaetous annelids from Japan, with descrip- 
tions of six new species.—Bulletin of the Na- 
tional Science Museum, Series A Zoology 1:5— 
I 

Levenstein, R. Y. 1977. A new genus and species of 
Polychaeta (Family Lumbrineridae) from the 
deep-water trenches of the North Pacific. Pp. 
189-198 in D. J. Reish & K. Fauchald, eds., 
Essays on the Polychaetous Annelids in Mem- 
ory of Dr. Olga Hartman. Allan Hancock Foun- 
dation, Los Angeles, 604 pp. 

McIntosh, W. C. 1903. Notes from the Gatty Marine 
Laboratory, St. Andrews, no. 25. 1. On the Eu- 
nicidae dredged by H.M.S. Porcupine in 1869 
and 1870. 2. On Canadian Eunicidae dredged 
by Dr. Whiteaves, of the Canadian Geological 
Survey, in 1871-1873. 3. On Norwegian Eu- 
nicidae collected by Canon Norman.—Annals 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


and Magazine of Natural History, Series 712: 
128-166. 

Miura, T. 1980. Lumbrinereidae (Annélides Polyche- 
tes) abyssaux récoltés au cours de campagnes 
du Centre Océanologique de Bretagne dans 
V’Atlantique et la Mediterranée—Bulletin du 
Muséum d’Histoire Naturelle, Paris, 2 section 
A 1019-1057. 

Monro, C. 1930. Polychaeta worms.—Discovery Re- 
ports 2:1—222. 

Moore, J. P. 1905. New species of Polychaeta from the 
north Pacific, chiefly from Alaskan waters.— 
Proceedings of the Academy of Natural Scienc- 
es of Philadelphia 57:525—554. 

. 1911. The polychaetous annelids dredged by 
the U.S.S. “Albatross” off the coast of South- 
ern California in 1904, 3. Euphrosynidae to 
Goniadidae.—Proceedings of the Academy of 
Natural Sciences of Philadelphia 63:234—-318. 

Orensanz, J. M. 1990. The eunicemorph polychaete 
annelids from Antarctic and Subantarctic Seas, 
with addenda to the Eunicemorpha of Argenti- 
na, Chile, New Zealand, Australia and the 
Southern Indian Ocean.—Antarctic Research 
Series 52:1—184. 

Oug, E. 1998. A new small species of Lumbrineris 
from Norway and Arctic waters, with comments 
on L. minuta (Théel, 1879) and L. vanhoeffeni 
(Michaelsen, 1898) (Polychaeta: Lumbrineri- 
dae).—Ophelia 49:147—-162. 

Théel, H. J. 1879. Les annelids polychétes der Mers 
de la Nouvelle-Zemble.—Kungliga Svenska 
Vetenskapsakademiens Handlingar 16:3—75. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):725—736. 2001. 


On the rotifer fauna of Bermuda, including notes on the associated 
meiofauna and the description of a new species of Encentrum 
(Rotifera: Ploima: Dicranophoridae) 


Martin V. Sgrensen 


Invertebrate Department, Zoological Museum, University of Copenhagen, Denmark. E-mail: 
mvsorensen @ zmuc.ku.dk 


Abstract.—The rotifer fauna of Bermuda was investigated by examining 
samples from 12 localities, including marine, brackish, and freshwater locali- 
ties. The majority of the 14 species that were identified are common cosmo- 
polites, but some also have a more limited distribution. No endemics were 
recorded. The dominant genus present in the samples, Lecane, was represented 
by 5 species. The relatively small total number of species is probably due to 
one or both of two factors: the lack of stable freshwater bodies, and Bermuda’s 
isolated geographic position. All but one of the recorded species are new to 
Bermuda. One species, Encentrum astridde, is new to science. The species has 
previously been found in Danish waters. Finding the new species in both Dan- 
ish and Bermudan waters suggests that it is distributed in at least the boreal 


and tropical West Atlantic regions. 


Few studies concerning rotifers from iso- 
lated oceanic islands have been accom- 
plished. However, to understand the zoo- 
geographic dynamics and dispersal poten- 
tial of rotifers it is important to know the 
distribution of these island-dwelling spe- 
cies. These include Galapagos (De Smet 
1989a, 1989b; Segers & Dumont 1993a), 
the Easter Island (Segers & Dumont 
1993a), the South Pacific islands (Russell 
1957), and the Azores (Green 1992). This 
study presents the first faunistic study on 
the rotifer fauna of the Bermuda archipel- 
ago. The localities studied include marine, 
brackish, and freshwater bodies. 

Bermuda is located in the Atlantic Ocean 
at about 32°N, 64°W, almost 1000 km SE 
of North Carolina, USA, which represents 
the nearest continental coast. The Bermuda 
seamount was first formed by volcanic 
eruptions about 100 mya, and re-erupted 35 
mya, covering an area much larger than the 
present area of Bermuda. Subsequent ero- 
sion, formation of coral reefs, a covering of 
limestone accreted by the wind-blown skel- 


etons of reef organisms, and the postglacial 
rise of the sea level formed the islands’ pre- 
sent appearance (Watson et al. 1965). 

On land, dune hills and limestone dom- 
inate Bermuda. There are no streams or riv- 
ers and only a few canals. A few lakes and 
ponds are present, but all are to some extent 
connected with the sea and therefore per- 
manently or temporarily brackish. 

Bermuda’s location in the Gulf Stream 
provides the islands a climate with hot sum- 
mers and mild winters. This enables the 
corals to form large reefs, and provide the 
calcareous coral sand that dominates the sea 
floor around Bermuda. This coral sand has 
proved to host a rich marine interstitial fau- 
na (Higgins 1982; Eibye-Jacobsen & Kris- 
tensen 1994: Sterrer 1998a, 1998b). For a 
more comprehensive review of the Ber- 
mudan fauna see Sterrer (1986). 


Materials and methods 


During a three-weeks stay at the Bermuda 
Aquarium Museum and Zoo (BAMZ), six- 


726 


teen localities were sampled for the purpose 
of collecting rotifers and gnathostomulids 
(Fig. 1). Seven marine, four brackish, and one 
freshwater station yielded rotifers (Tables 1, 
2). 

Intertidal samples were dug up with a 
shovel, while subtidal samples were taken 
by snorkeling. In both cases the upper 10 
cm of the sediment was collected, placed in 
buckets, and returned to the laboratory for 
processing. Meiofaunal organisms were ex- 
tracted from the sediment by using an iso- 
tonic solution of magnesium sulfate to 
anaesthetize the animals. Approximately 1 
liter sediment was transferred to a conical 
plastic bottle and a corresponding amount 
of magnesium sulfate solution was added. 
After a ten minutes incubation period the 
containers were agitated vigorously, and af- 
ter a brief period settling the supernatant 
was decanted through a 30 4m mesh sieve. 
This procedure was repeated once. Samples 
taken from detritus, algae and plants were 
Squeezed and concentrated in a 30 wm mesh 
sieve. 

All samples were sorted using a Wild 
M420 dissection microscope and a Wild 
M20 microscope. Rotifers were identified 
and drawn using a Wild M20 compound 
microscope with camera lucida. Trophi 
were isolated by dissolving the animals us- 
ing dilute sodium hypochlorite and then 
prepared for SEM and light microscopy (De 
Smet 1998). Trophi for light microscopy 
were mounted in a modified Faure’s solu- 
tion. Trophi for SEM were examined and 
photographed with a JEOL JSM-840 micro- 
scope. All material is deposited in the Zoo- 
logical Museum, University of Copenhagen 
(ZMUC). 


Results 


Sixteen localities investigated yielded a 
total of 14 identified rotifer species, two 
forms of the same species, a number of un- 
identified bdelloid species from the genera 
Philodina and Rotaria, and one dead spec- 
imen of Monommata which could not be 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


identified. All recorded species are listed in 
Table 3. 

Five species, of the genera Colurella, En- 
centrum, and Proales, were recorded from 
the marine localities. One of these, Encen- 
trum astridae new species, is new to sci- 
ence. The brackish water localities yielded 
eight species, distributed among six genera. 
Furthermore, some unidentified Rotaria 
were recorded. Two species, Colurella un- 
cinata uncinata and Proales similis, oc- 
curred both at marine and brackish locali- 
ties. The single freshwater locality pro- 
duced three identified species, one uniden- 
tified Monommata and some Philodina spp. 
Colurella uncinata was recorded from the 
freshwater locality as well as from a brack- 
ish and a marine locality. In the freshwater 
locality the species was represented by the 
form C. uncinata bicuspitata. 

The remaining interstitial fauna at the 
marine localities was only investigated su- 
perficially. However, it was noted that the 
samples contained a rich meiofauna. Most 
samples contained several species of ma- 
crodasyid gastrotrichs, including the easily 
recognized genus Urodasys. Nematodes 
were numerous, including different des- 
moscolecids. Also several interstitial poly- 
chaetes, including the families Dorvilleidae 
and Syllidae were represented. Five species 
of Gnathostomulida, Haplognathia rosea 
(Sterrer, 1969), Tenuignathia rikerae Ster- 
rer, 1976, Problognathia minima Sterrer & 
Farris, 1975, Gnathostomula peregrina Kir- 
steuer, 1964, and Austrognathia christianae 
Farris, 1977, were recorded. Gnathostomula 
peregrina was the most abundant and was 
present at stations 3—7. Two species of Ki- 
norhyncha, Antygomonas cf. oreas Bauer- 
Nebelsick, 1996 and Echinoderes bermu- 
densis Higgins, 1982, were recorded from 
station 7 and 4, respectively. Except for the 
arrangement of the lateral spines on seg- 
ment 11 the Antygomonas species fits per- 
fectly the description given by Bauer-Ne- 
belsick (1996). In the specimens described 
by Bauer-Nebelsick (1996) the cuspidate 
spines are located laterally to the acicular 


VOLUME 114, NUMBER 3 


A 


Harrington 


Sound 
SS) 
Loc. 5 


Loc. 1-2 
Loc. 6-7 


Spittal Park 
Pond 
Loc. 12 EE —Loe. 13 
Loc. 11 


| Loc. 10 
Loc. 14 


Loc. 15 Loc. 4 
@ 


~ 


Castle 
Harbour 


VT 


Fig. 1. Map showing location of sampling sites on Bermuda. Localities 2, 13, 14, and 15 yielded no rolifers 
and are omitted in the following tables, thus Loc. 1 = st. 1, Loc. 2 = st. 2, Loc. 4 = st. 3, Loc. 5 = st. 4, etc., 


and Loc. 16 = st. 12. 


728 


Table 1.—Data on marine localities. Salinity on all stations ca. 33%o. 


io 


St. No. 


Mouth of Flatts In- 


Gibbon Bay 


Gibbon Bay 


BAMZ docks Tuckers Town At Trunk Island 


Flatts Inlet 


Name 


let 


32°19'18"N 


Cove 
32°19'49"N 


64°41'30"W 


32°19'16"N 
64°44'3 1" W 


32°19'16"N 


32°19'51"N 


32°19'21°N 


32°19'19"N 


Position 


64°43'32"W 


64°44'3 1" W 


64°43'39"W 


64°44'10"W 


64°44'17"W 


15 Sep 2000 


3m 


BIC 


17 Sep 2000 


15 Sep 2000 


0.5 m 


7 Sep 2000 7 Sep 2000 


6 Sep 2000 
2m 
28°C 
from fire Subtidal, 


6 Sep 2000 


1m 


28.4°C 


Coll. Date 


Depth 
Temp. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


29.2°C 
from fine Intertidal from fine, Subtidal, 


28.2°C 


DRE 


28.3°C 
from red Intertidal, from me- 


from me- 
clean coral 


from me- 


Tedania algae on reef and dium coral sand w. dium coral sand w. coral sand w. detri- clean coral sand 


Remarks 


Subtidal, 


Subtidal, 


Subtidal, 


dium, 
sand 


sponges, 
ignis 


tus 


detritus 


detritus 


docks 


spines, while the position of these have 
shifted in the Bermuda specimen, so the 
acicular spines on segment 11 are most lat- 
eral. Antygomonas oreas was described 
from coral sand at relatively deep water 
(500—600 m) in the Pacific Ocean (Bauer- 
Nebelsick, 1996). This is the first reported 
record of the species since then. 


Family Dicranophoridae Harring, 1913 
Genus Encentrum Ehrenberg, 1838 
Encentrum astridae, new species 
Figs. 2—3 


Type material.—All type material was 
obtained from samples taken on 15 Septem- 
ber 2000 at station 5, Gibbon Bay close to 
Flatts Inlet, Bermuda (Table 1). Position: 
32°19'16"N, 064°44'31"N. Holotype: Adult 
female, mounted in glycerol (ZMUC ROT- 
223). Paratypes: 9 adult females, mounted 
in glycerol (ZMUC ROT-224 to ROT-232); 
2 isolated trophi mounted for LM (ZMUC 
ROT-233 and ROT-234); 3 isolated trophi 
from adults mounted for SEM (ZMUC 
ROT-235 to ROT-237); 1 isolated trophus 
from a juvenile mounted for SEM (ZMUC 
ROT-238). All types are stored at ZMUC. 

Other material.—Further material was 
obtained 7 July 1999, at Prestebugten, Hir- 
sholmene, Denmark, a small group of is- 
lands in the northern Kattegat off Freder- 
ikshavn, Denmark. Samples were taken in 
the tidal zone from well-sorted medium 
sized quartz sand covered with sulfur bac- 
teria. Position: 57°29'17"N, 010°37;'29’E. 2 
isolated trophi mounted for SEM (ZMUC 
ROT-244 and ROT-245) and stored at 
ZMUC. 

Diagnosis.—Large animals (ca. 350 wm 
long), trunk with longitudinal folds (Fig. 
2A-—B). Foot ventrally displaced (Fig. 2B). 
Posterior part of stomach covered by yel- 
lowish to brownish glandular syncytium 
with brown nuclei (Fig. 2A—B). Trophi 
large, forcipate. Rami large with sharp api- 
cal rami teeth, and a pair of smaller, hook- 
shaped preuncinal teeth. Unci very long, al- 
most as long as rami length, with sharp in- 


VOLUME 114, NUMBER 3 


Table 2.—Data on brackish and fresh water localities. All samples taken just below the water surface. 


11 


10 


St. No. 


Mangrove Lake 
32°19'26"N 
64°42'38"W 


Pond in Zoo, BAMZ 


32°19'20"N 
64°44'18"W 


Spittal Pond, west 
32°18'32"N 
64°43'39"W 


Spittal Pond, east 
32°18'39"N 
64°43'29"W 


Bird Sanctuary Pond 


32°18'34’N 
64°43'33"W 


Name 


Position 


15 Sep 2000 


O0%o 


14 Sep 2000 
—12%o 
29.6°C 


12 Sep 2000 
12-18% 
25.6°C 


12 Sep 2000 


3-8%o 
26.4°C 


7 Sep 2000 
10-15%c 


ae 


Coll. Date 
Salinity 
Temp. 


25.1°C 


algae 


From plankton, 


and plants 


From detritus and de- 


graded plants 


From detritus and de- 


graded plants 


From red bacteria mats 
and degraded plants 


From algae on man- 


grove roots 


Remarks 


T29) 


ward-curved teeth. Supramanubria with 
long curved extensions, meeting each other 
in center of trophi (Figs. 2C, 3). 

Description.—Body elongate, fusiform. 
Head medium size; rostrum short; corona 
slightly oblique, ventrally. Brain large, sac- 
cate, extending into neck; subcerebral 
glands elongate; no light-refracting ele- 
ments; retrocerebral sac absent; eyespots 
absent (Fig. 2A—B). Neck short. Trunk with 
three distinct tranverse folds; posterior 
pseudosegment with broad tail. Dorsal and 
lateral sides of trunk with irregularly dis- 
tributed longitudinal folds. Mastax large, 
with pair of salivary glands. Proventriculus 
present. Stomach surrounded by yellowish 
to brownish glandular syncytium with 
brown nuclei; number of nuclei ranging be- 
tween 12 and 19 (Fig. 2A—B). Gastric 
glands elongate, close to stomach. Vitellar- 
ium large, elongate. Foot long, conical, re- 
tractable, ventrally displaced (Fig. 2A—B). 
Toes close-set, almost parallel sided, taper- 
ing distally to tips. Pedal glands elongate, 
narrow (Fig. 2A—B). 

Trophi forcipate, large, elongate, slender 
(Figs. 2C, 3). Rami expanded %4 from prox- 
imal ends; outer margins slightly concave 
medially, slightly diverging distally (Figs. 
2C, 3A—C). Median opening wedge-shaped. 
Each ramus tip terminating dorsally in a 
sharp, slightly curved, inwardly projecting 
apical ramus tooth (Figs. 2C, 3A, C); ven- 
trally, at bases of apical rami teeth, a small, 
stout, slightly hook-shaped, inwardly pro- 
jecting preuncinal tooth (Figs. 2C, 3B, D). 
Scapus long, narrow, extending into ante- 
rior part of ramus (Fig. 2C); fenestra of sca- 
pus small, opening basally on dorsal side of 
each ramus (Fig. 3C). Bulla small, almost 
rectangular (Fig. 2C); fenestra of bulla 
small, opening caudally on ramus (Fig. 3D, 
F). Fulcrum rod-shaped in dorsal view 
(Figs. 2C, 3A—C), gradually tapering to- 
wards truncate tip in lateral view (Fig. 3D). 
Unci very long, almost 9/10 of rami length; 
shaft long, rod-shaped; teeth long, sharp, in- 
wardly curved, with small dorsal apophysis 
at bases (Figs. 2C, 3). Manubria gradually 


730 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


curved inwardly towards distal ends (Figs. 
2C, 3); proximal ends with anterior and me- 
dian chamber retained. Anterior chamber 
present in dorsally pointed extension; me- 
dian chamber with small fenestra ventroap- 
ically on manubrium. Intramallei small, tri- 
angular, attached to ventral side of supra- 
manubria (Figs. 2C, 3B). Supramanubria 
. very large, with stout bases tapering into 
AA long extensions with curved terminals; ter- 
minals meet each other in center of trophi 
(Figs. 2C, 3A—C). 
is y y Juveniles about 2/3 of adult size; very 
hyaline, no coloration of any organs yet; 
trophi as in adults (Fig. 3B). No males were 
a od vaw ».| found. 
Measurements of adult females: Body 
length 308-388 wm; toes 26—31 wm; trophi 
6 Sd 3d 38 wm; rami 22 pm; fulcrum 10 pm; unci 
19 wm; manubria 24—26 pm; intramallei 3 
zm; Supramanubria 10 wm. 
wn < > >< Etymology.—The species is named after 
my daughter Astrid. 
Remarks.—The presence of relatively 
+ x x x simple forcipate trophi with intramallei and 
supramanubria, and a long fulcrum, com- 
bined with the absence of teeth on inner 
ca a margins of rami and single-toothed unci 
clearly place Encentrum astridae new spe- 
cies in the genus Encentrum (see De Smet 
x rs IATA 1997). The presence of elongate rami with 
very long scapus, dorsal incurved apical ra- 
mus teeth and ventral preuncinal teeth set 
at right angles to rami place the species in 
subgenus Encentrum s. str. Ehrenberg, 1838 
(see De Smet 1997). 


10 
x 


ass 


A 

le) 

n 

i= 

ie) 

3 

Z 

S . . 

g a a The species can hardly be confused with 

Py ~% es ~ | other known species due to the presence of 
ing) a Zao tom n . . . . . 

tS 2 © iS e a = a large trophi with very long unci with in- 

B= Fee) SN 20 <.| wardly pointed teeth, and long supramanu- 
O & op OS) a 2 3 : : 

a se e 9 Eo J ood Bs 5 brial extensions reaching each other (Figs. 

io) é : He) = Qa to} aes 9) 

ale |eSRess ze e225 31 2C, 3A-C). 

Ss = [30 I HDonwm. S - o 

S |2ig8 Se es eeees eases 2 The species was found at five of the in- 

to = & (So) & Ls OD = «AD OQ 

5 Seaggeess 227588 2| vestigated marine localities (see Table 3), 

oO YS PAs es Se) i 2) 9 é i 

fe) ene S36 S: Ss g © S S => &._.-2]| and is probably common in the psammon 
BS) S SS ps) SX , 8 & Ss = aq "= : a € 

I 2s g S) Sane SS 86 5 8 S 2 =} around Bermuda. It was found in intertidal 
ESSSPSSSTESSEse ss 

a) 8 < = SR REE S = 2ESSE = o| as well as subtidal psammon, but was most 

2 Ss oS Ss S&S 5-8 S 2S 8 ws 5 P 

= SESSCSSESSSES SoS S | abundant in the subtidal. It occurred on 
RS ES) oo ao oo SS me ass aS = 
SOOUUURReIG454NS2a] depths ranging from 0 to 3 meter. In June 


VOLUME 114, NUMBER 3 


733 


Fig. 2. Encentrum astridae new species. A. Female habitus, dorsal view. B. Female habitus, lateral view. C. 
Trophi, ventral view. Abbreviations: ar, apical ramus tooth; br, brain; bu, bulla; fu, fulcrum; ga, gastric glands; 
gl, glandular syncytium; im, intramalleus; ma, manubrium; mx, mastax; pg, pedal glanda; pr, proventriculus; pu, 
preuncinal tooth; ra, ramus; sc, scapus; sg, subcerebral glands; sm, supramanubrium; st, stomach; un, uncus; vi, 


vitellarium. 


1999 it was found in psammon samples 
from Danish waters. The species was found 
during meiofauna collections on Hirshol- 
mene, a small group of islands in the north- 
erm part of Kattegat, off Frederikshavn, 
Denmark, but was not described then due 
to insufficient material. This distribution 
suggests that the species may be found in 
well-sorted sandy sediments at shallow wa- 
ters in most of the Northern Atlantic. 


Discussion 


No systematic study has been made of 
the Bermudan rotifer fauna before. Only 
four species were previously recorded, and 
none of these were identified to more than 
genus level (von Bodungen et al. 1982; 
Sterrer 1986, 1998a). von Bodungen et al. 
(1982) recorded Trichocerca sp. from Ham- 
ilton Harbour and Sterrer (1986) reports 
Encentrum sp., Lindia sp., and Synchaeta 


sp. from different localities, and illustrates 
the recorded species (Sterrer 1986: plate 
69). Encentrum sp. is probably identical to 
Encentrum tectipes (Fig. 4A—B). Lindia sp. 
is identical to L. tecusa Harring & Myers, 
1922, due its presence in sublittoral psam- 
mon. This identity was confirmed during 
my stay at BAMZ, where I had the oppor- 
tunity to investigate unpublished LM pic- 
tures of the species. The species was not 
recorded during this study, and Sterrer 
(1986) refers to it as “rare”. Neither Syn- 
chaeta sp. nor Trichocerca sp. were record- 
ed in this study. Both species were found 
in planktonic samples (von Bodungen et al. 
1982; Sterrer 1986), and since the marine 
samples in this study only were taken from 
psammon or phyton, it explains why these 
were not recorded. Based solely on the il- 
lustrations, the Synchaeta species reported 
by Sterrer (1986) cannot be identified. von 


732 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. 


Bodungen et al. (1982) do not illustrate the 
recorded Trichocerca sp, but it might be 
identical to T. marina (Daday, 1890), which 
is the only known marine planktonic 77i- 
chocerca. 

Lecane was the dominating species and 
accounted for 31% of the total amount of 
identified species. Though the material in 
this study is rather small, it confirms the 
pattern that Lecane displays high species 
diversity in tropical and subtropical regions 
(see Harring 1914; De Smet 1988; Segers 
& Dumont 1993a, 1993b, 1995; Segers & 


Encentrum astridae new species. SEM photos of trophi. A is a Danish specimen, B—F is Bermudan 
specimens. A. Dorsal view. B. Ventral view, from juvenile specimen. C. Dorsal view, rami opened. D. Lateral 
view. E. Frontal view. EF Caudal view. Abbreviations: ar, apical ramus tooth; bu, bulla; fu, fulcrum; im, intra- 
malleus; ma, manubrium; pu, preuncinal tooth; ra, ramus; sc, scapus; sm, supramanubrium; un, uncus. 


De Meester 1994; Segers & Sanoamuang 
1994; Janetzky et al. 1995; Sanoamuang 
1998; Samraoui et al. 1998). 

A few specimens of Cephalodella forfi- 
cata (Fig. 4E—F) were recorded from one of 
the brackish water localities (Table 3). All 
specimens were juveniles, and the gastric 
glands were only light red. The species is 
considered cosmopolitan, but is mainly 
found in freshwater (Koste 1978; Nogrady 
& Pourriot 1995). 

Proales similis (Fig. 4C—D) was found at 
brackish as well as marine localities (Table 


VOLUME 114, NUMBER 3 


733 


Fig. 4. SEM photos of trophi. A. Encentrum tectipes, dorsal view. B. E. tectipes, ventral view. C. Proales 
similis, ventral view. D. P. similis, rami, dorsal view. E. Cephalodella forficata, ventral view. E C. forficata, 
lateral view. Abbreviations: al, alula; bf, basifenestra; bl, basal lamella; fe, fulcral extension; fu, fulcrum; im, 
intramalleus; ma, manubrium; ra, ramus; sc, scapus; sf, subbasifenestra; sm, supramanubrium; un, uncus. 


3). The species is recognized by its fusi- 
form trunk and offset foot with one pseu- 
dosegment and distinct wrinkles. The apical 
rami tips were rather short (Fig. 4D) in the 
recorded specimens, and not extended as in 
the specimens pictured by De Smet (1996). 
The species has been recorded from most 
parts of the world and is found in marine 
waters as well as inland saline or brackish 
ponds (De Smet 1996). 

Beside Encentrum astridae new species, 
another dicranophorid, E. tectipes, was re- 


corded from the marine localities. The spe- 
cies is easily recognized by its robust and 
compact trophi (Fig. 4A—B), and the pres- 
ence of yellowish glandular tissue around 
the posterior part of the stomach, also de- 
scribed by Remane (1949). Like many other 
marine, interstitial dicranophorids, the spe- 
cies has formerly only been recorded from 
the northwestern Europe (De Smet 1997), 
but finding the species on Bermuda sug- 
gests that it is much more widely distrib- 
uted. As a matter of fact, the known distri- 


734 


butional patterns of many marine, intersti- 
tial rotifers with an apparently limited dis- 
tribution are probably mostly a reflection of 
our insufficient knowledge of species from 
such habitats, rather than a picture of their 
actual distribution. Recently, E. tenuidigi- 
tatum De Smet, 2000, described from tidal 
psammon in Belgium, was recorded from 
Greenland (Funch & Sgrensen 2001), and 
E. porsildi Sgrensen, 1998 described from 
Disko Island, Greenland, was recorded 
from Denmark (Sgrensen, in press). I find 
it very likely that the interstitial habitat in 
clean, well-sorted sand offers a stable en- 
vironment, and since marine environments 
generally are more climatologically stable 
than terrestrial, it enables the psammobion- 
tic species to be widely distributed and 
makes factors as climate less important. 
This is also suggested by the distribution of 
E. astridae new species. The study of the 
marine psammobiontic rotifers has often 
been neglected compared to the more spe- 
cies-diverse freshwater habitats, but if more 
attention were paid to this special habitat, 
our knowledge would certainly be increased 
greatly. 

The fauna on isolated oceanic islands can 
theoretically be expected to have low spe- 
cies diversity and a relatively high propor- 
tion of endemism (Segers & Dumont 
1993a). A total of 14 identifiable species, 
as recorded in this study, must be consid- 
ered a low number of species compared to 
the number of investigated localities. How- 
ever, it is most likely that this is caused by 
the lack of stable freshwater bodies rather 
than isolation by distance. This does not 
overrule that isolation also plays a role. Of 
the total number of species, 65% are cos- 
mopolites while three, L. grandis, L. has- 
tata, and L. punctata, have a tropical/sub- 
tropical distribution. Two species, E. astri- 
dae new species and E. tectipes, have at 
least a boreal and tropical West Atlantic dis- 
tribution. No endemics were recorded. The 
dominance of cosmopolites can probably be 
explained by the fact that migrations of 
such species are more likely to occur than 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


migration of species with a more limited 
distribution. 


Acknowledgments 


I am indebted to W. Sterrer for his help, 
support and suggestions to the manuscript, 
and to Bermuda Zoological Society and the 
staff at BAMZ for their hospitality. I thank 
the reviewers H. Segers and R. L. Wallace 
for their suggestions to the manuscript and 
M. E. Petersen for correcting the language. 
The study was funded by Bermuda Zoolog- 
ical Society and the University of Copen- 
hagen. This paper is contribution #43 from 
the Bermuda Biodiversity Project (BBP), 
Bermuda Aquarium, Natural History Mu- 
seum and Zoo. 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):737—745. 2001. 


New Unguiphora (Platyhelminthes: Proseriata) from India 


Marco Curini-Galletti, Gavino Oggiano, and Marco Casu 


Dipartimento di Zoologia ed Antropologia Biologica, Universita di Sassari, via Muroni 25, 
I-07100 Sassari, Italy 


Abstract.—Three new species of Unguiphora are described from Goa (India). 
Two species belong to the genus Nematoplana (N. indica and N. calamus). 
They are distinguished from the known species of the genus on the basis of 
the morphology of sclerotized structures. The new genus Alloeostyliphora is 
established for the third species (A. mirabilis), based on the unique features of 
its copulatory apparatus, with two symmetrical copulatory organs, and a few, 


non-glandular accessory stylets. 


The Unguiphora is the smallest surborder 
of the Proseriata (Platyhelminthes). At pre- 
sent, the taxon is comprised of 5 genera and 
34 species. In general, very few species are 
known from any given biogeographical 
area (Curini-Galletti & Martens 1991, 
1992). However, the recent finding of an 
exceedingly rich unguiphorid fauna in east- 
ern Australia raises the question of our ac- 
tual understanding of the diversity of the 
taxon in extra-boreal areas (Curini-Galletti 
1998, Curini-Galletti et al. 2001). The pre- 
sent contribution deals with three ungui- 
phorid species found in a small sediment 
sample from Goa, India. 


Materials and Methods 


The sample (about one-half liter of sed- 
iment) was collected in intertidal pockets of 
clean medium-fine sand among rocks, in 
front of Fort Aguada Beach Hotel, Aguada, 
Goa (India) (May 1993). The animals were 
extracted from the sediment in the labora- 
tory with the MgCl, decantation technique 
(Martens 1984). Preservation and histolog- 
ical techniques routinely adopted for Pro- 
seriata were used (see Martens et al. 1989); 
whole mounts were made with polyvinyl- 
lactophenol or Faure. Karyological tech- 
niques are described in Curini-Galletti et al. 
(1989). Idiograms (Figs. 2E, 3G) are based 


on karyometrical data presented in the kar- 
yotype formula: haploid genome absolute 
length in wm, relative length and centro- 
mere index of each chromosome; chromo- 
some nomenclature between parentheses (m 
= metacentric; sm = submetacentric). Type 
material is deposited in the collections of 
the Queensland Museum, S. Brisbane, Aus- 
tralia (QM). 


Family Nematoplanidae Meixner, 1938 
Genus Nematoplana Meixner, 1938 
Nematoplana indica, new species 
Figs. 1, 4A 


Material examined.—Holotype, India, 
Goa, Aguada: whole mount (lactophenol) 
(QM-G211833). 

Etymology.—Named after the geograph- 
ical area, from which the species is record- 
ed. 

Description.—The holotype is an adult 
worm, about 4.2 mm long in fixed condi- 
tion, without pigment or pigmented eye- 
spots. Anterior end elongate, provided lat- 
erally and terminally with sensory bristles. 
The pharynx, short and collar shaped, is lo- 
cated in the posterior fifth of the body. 

Male genital organs: with numerous tes- 
tes irregularly arranged between vitellaria 
in front of the pharynx. The copulatory or- 
gan consists of an elongate, muscular bulb 


738 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 


Nematoplana indica, new species. (A) general organization of live animal; (B) stylet (from holotype); 
(C) detail of the post-pharyngeal area. Abbreviations in figures: ast—accessory stylet; br—brain; bl—bulb; co 


copulatory organ; e—eye-spot; fd—female duct; fg—female glands; fp—female pore; fsv—fused seminal ves- 


icles; g—gut; ma—male antrum; o—oocyte; ph 
stylet; t—testis; vi—vitellaria. 


(about 110 pm long), provided with a cop- 
ulatory stylet, and two seminal vesicles, 
which enter the bulb proximally. The sem- 
inal vesicles, which have a very thin lining, 
are fused caudally. The stylet is funnel 
shaped, 48 wm long, with a broad, mark- 
edly oblique, proximal opening and a nar- 
row distal opening (Fig. 1B). Its slender 
base is about 30 wm long; its maximum di- 
ameter (about 11 jm) is attained just below 
the apophysis, which is about 14 pm long 
and 4 wm wide. The apophysis is nearly 
perpendicular to the main axis of the stylet. 
Numerous, thick longitudinal muscles, most 
of which are connected basally with the 
musculature surrounding the bulb, are at- 
tached to the apophysis. The distal end of 
the stylet narrows above the apophysis into 
a nearly straight tube, about 0.33 of the to- 


pharynx; phg—pharyngeal glands; sv—seminal vesicle; st— 


tal length of the stylet. It is provided with 
a long, slightly recurve, distal tip (about 20 
wm long), perpendicular to the main axis of 
the stylet. 

Female genital organs: the specimen ex- 
amined possessed one mature oocyte, me- 
dially in front of the pharynx. Vitellaria ex- 
tend from posterior to the brain to in front 
of the ovaries, and from posterior to the 
pharynx to the level of the copulatory bulb. 
The oviducts fuse posterior to the bulb into 
a short common female duct, which opens 
to the outside through a female pore, me- 
dially between the seminal vesicles. 

Remarks.—The distinctly claw-shaped 
stylet, with an elongate base, a slightly re- 
curve distal tip, and a long apophysis, is 
comparable to that of the N. coelogynopo- 
roides species group (which includes N. 


VOLUME 114, NUMBER 3 


coelogynoporoides Meixner, 1938, N. cil- 
iovesiculae Tajika, 1979, N. riegeri Curini- 
Galletti & Martens, 1992 and N. cannoni 
Curini-Galletti, Oggiano & Casu, 2001) (cf. 
Curini-Gallettt & Martens 1992, Curini- 
Galletti et al. 2001). The insertion of the 
apophysis with the axis of the stylet differs; 
it is perpendicular in N. indica and oblique 
(at an angle of about 45°) in the species 
listed above. Furthermore, the stylet of WN. 
indica is straighter above the apophysis, 
with a much longer distal tip, that is more 
clearly orthogonal to the main axis, than 
any of the species listed above. Among the 
Indo-Pacific members of the N. coelogyn- 
oporoides group, the Japanese N. ciliovesi- 
culae has a more recurve, markedly larger 
(about 130 wm long) stylet, with a very 
short distal tip and a relatively very short, 
obtuse apophysis. Nematoplana cannoni 
from eastern Australia is similar to N. 
indica for the size of the stylet. Further- 
more, both species share the presence of 
unpaired oocytes, and of an elongate mus- 
cular bulb. However, in N. cannoni the por- 
tion of the stylet above the apophysis is re- 
curved and comparatively longer and 
broader than in N. indica. Furthermore, its 
stylet is provided with a much shorter distal 
tip; the apophysis is oblique, narrow, and 
angled distally; the basis is broader proxi- 
mally. Nematoplana cannoni has pigmented 
eye-spots, enclosed within the brain cap- 
sule. 


Nematoplana calamus, new species 
Figs. 2, 4C, D 


Material examined.—Holotype: India, 
Goa, Aguada: whole mount (lactophenol): 
QM-G211834. Paratype: one karyological 
Slide made permanent with Faure (QM- 
G211835), same data as holotype. 

Etymology.—The name refers to the 
shape of the stylet, which resembles an old- 
fashioned pen-nib (lat. calamus, a noun 
used as an apposition). 

Description.—Animals small: the holo- 
type is an adult worm, about 1.2 mm long 


739 


in fixed condition. Without pigment or pig- 
mented eye-spots. Anterior end elongate, 
provided laterally and terminally with sen- 
sory bristles. The short, collar-shaped phar- 
ynx is located in the posterior sixth of the 
body. 

Male genital organs: with a few testes ir- 
regularly arranged among vitellaria, in front 
of the pharynx. The copulatory organ con- 
sists of a single seminal vesicle, and a bulb 
provided distally with a stylet. The seminal 
vesicle is broadly elongate, and provided 
with a coating of circular musculature. The 
seminal vesicle enters the bulb at its prox- 
imal base. The ovoid bulb is about 20 wm 
long, and is provided with numerous pros- 
tatic glands, some of which have their cell 
bodies outside the bulb itself. The stylet 
(Figs. 2B, C, 4C, D), about 18 wm long, is 
thin and diaphanous. Its base (about 11 wm 
wide) is gutter-shaped, widely open proxi- 
mally and ventrally, with a marked notch at 
its dorsal side. The ventral sides are nearly 
straight, and are distinctly angled distally. 
The stylet is provided distally with a re- 
curve, convex distal spike, about 10 wm 
long. In living specimens, the stylet ap- 
peared distinctly claw-shaped. Presumably, 
it acts functionally as a funnel, with the dis- 
tal opening located beneath the dorsal 
spike, and the proximal opening formed by 
the adjoining of the ventral flaps. 

Female genital organs: mature oocytes 
were not present. Vitellaria extend from be- 
hind the brain to in front of the pharyx. 
Female pore, surrounded by female glands, 
posterior to the male pore. 

Karyotype: most chromosomes are meta- 
centric (n = 6). Karyotype formula: 10 wm; 
I: 22.49, 46.36 (m); II: 18.7, 40.92 (m); III: 
15.98, 44.16 (m); IV: 15.23, 29.54 (sm); V: 
15.1, 43.15 (m); VI: 12.47, 43.56 (m). 

Remarks.—The stylet of Nematoplana 
calamus is peculiar. In no other unguiphorid 
species is it gutter-shaped, gaping ventrally, 
and with a distal opening located beneath a 
dorsal spike. The presence of a single sem- 
inal vesicle is shared with N. martensi Cur- 
ini-Galletti, Oggiano & Casu, 2001, N. ha- 


740 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. 


Nematoplana calamus, new species. (A) general organization of living animal; (B, C) stylet: (B) 


dorsal view (from holotype); (C) lateral view (from paratype); (D) detail of post-pharyngeal area; (E) idiogram. 


See Fig. 1 for explanation of abbreviations. 


mata Curini-Galletti, Oggiano & Casu, 
2001, N. cyclops Curini-Galletti, Oggiano 
& Casu, 2001, and N. pullolineata Tajika, 
1979 (Curini-Galletti et al. 2001, also for 
discussion of the problematical N. naia 
Marcus, 1949). The single-vesicle condition 
has been considered as a secondarily de- 
rived character status for the genus Nema- 
toplana, resulting from the total fusion of 
the two partially fused vesicles found in the 
other species (Curini-Galletti & Martens 
1992). This condition is not reported for 
any other Unguiphora. The Nematoplana 
species cited above share a further derived 
condition, 1.e., the presence of a cylindrical 
stylet, not provided with an apophysis, and 
have been considered as constituting a 
monophyletic group within the genus Ne- 
matoplana (Curini-Galletti et al. 2001). Ne- 
matoplana calamus lacks any obvious 


apophysis; the structure of its stylet is, how- 
ever, SO apomorphic that, at the moment, it 
precludes the inclusion of N. calamus in 
any evolutionary line known within the Un- 
guiphora. The inclusion of N. calamus (as 
well as of the species listed immediately 
above, see Curini-Galletti et al., 2001) with- 
in the genus Nematoplana should be con- 
sidered as conservative, until more data on 
polarization and distribution of characters 
are available, and a more significant frac- 
tion of the world species is described. 


Alloeostyliphora, new genus 


Diagnosis.—A new genus of Unguiphora 
with two symmetrical copulatory organs, 
provided with reduced bulbs. With acces- 
sory (replacement?) stylets, not connected 
to glandular structures. With a large male 
antrum anterior to the copulatory organs. 


VOLUME 114, NUMBER 3 


Type species.—Alloeostyliphora mirabi- 
lis, new species (by monotypy). 

Etymology.—The generic name is coined 
from alloeo, latinized from Greek alloios = 
different + styliphora from the generic 
name Polystyliphora Ax, 1958. The generic 
name is feminine. . 


Alloeostyliphora mirabilis new species 
Figs. 3, 4B 


Material examined.—Holotype: India, 
Goa, Aguada: whole mount (lactophenol): 
QM-G211836. Paratype: one specimen sag- 
ittally sectioned (QM-G211837), same data 
as holotype. One immature studied karyo- 
logically. 

Etymology.—mirabilis (‘admirable’) re- 
fers to the striking features of the new spe- 
cies. 

Description.—Animals very small for 
the group: the holotype is about 0.8 mm 
long. With two pigmented eye-spots located 
within the brain capsule. Anterior end elon- 
gate, provided laterally and terminally with 
sensory bristles. Subepidermal longitudinal 
musculature well developed on the ventral 
side. Epithelium entirely ciliated (cilia 
length about 1 wm), with non-insunk nuclei. 
The short, collar-shaped pharynx is located 
in the posterior fifth of the body. Its epithe- 
lium has insunk nuclei, and is ciliated ex- 
- cept for a small area at the distal tip, where 
a few pharyngeal glands discharge. The cell 
bodies of these glands are located outside 
the pharynx itself. Pharyngeal cilia about 
1.5 wm long. No esophageal area could be 
seen. 

Male genital organs: with very few (one 
to two), large testes among vitellaria well 
in front of the pharynx. With two symmet- 
rically paired copulatory organs, posterior 
to the pharynx. Each consists of an elongate 
seminal vesicle connected to a copulatory 
stylet. One of the copulatory organs of the 
holotype had a proximally bifurcated sem- 
inal vesicle; each end was connected to a 
copulatory stylet. A variable number of ac- 
cessory stylets (1-3) were observed, lying 


741 


very close to the copulatory stylets. In liv- 
ing animals, they were arranged in a row, 
just caudal to the copulatory stylets. These 
accessory stylets are not connected to the 
seminal vesicle, nor, apparently, to any 
glandular structure. The copulatory and ac- 
cessory Stylets are tubular structures, iden- 
tical in morphology. The proximal opening 
is about 10 wm wide. The dorsal side of the 
stylet is markedly oblique, while the ventral 
side is straighter, about 7 ~m long. The dis- 
tal opening is about 4 ym in diameter. The 
stylet is provided with a straight or slightly 
recurved tip, about 7 wm long. In sections, 
a very small glandular area is seen, at the 
connection of the seminal vesicle with the 
copulatory stylet; this may be considered as 
homologous to the bulb of other species of 
Unguiphora. Each copulatory stylet pro- 
trudes into a small antrum, which opens 
into a broad male antrum. 

Female genital organs: with only one oo- 
cyte, medially in front of the pharynx. Vi- 
tellaria extend from behind the brain to in 
front of the pharynx. Female pore behind 
the male pore. 

Karyotype: chromosome | is appreciably 
larger than the other pairs (n = 5). Chro- 
mosomes are meta- or submetacentric. Kar- 
yotype formula: 6.5 wm; I: 29.33, 44.92 
(m); HI: 19.58, 46.73 (m); III: 19.16, 46.19 
(m); IV: 17.44, 36.94 (sm); V: 14.48, 36.62 
(sm). 

Remarks.—Alloeostyliphora mirabilis 
presents a remarkable set of features, which 
justifies the establishment of a new genus. 
Among the most striking is the fact that no 
other Proseriata Unguiphora has two func- 
tional copulatory organs. Among Proseriata 
Lithophora, Monotoplana diorchis Meixner, 
1938 has a continuous production of cop- 
ulatory organs, which are arranged into an 
irregular row (Meixner 1938). In M. dior- 
chis, however, only the distalmost copula- 
tory organ is functional at any time and ap- 
pears to be progressively replaced by a new 
one. In A. mirabilis, the two organs are 
symmetrical, and both appear to be func- 
tional at the same time. The presence of two 


742 


Fig. 3. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


phg 


ma 


20 


Alloeostyliphora mirabilis, new genus, new species. (A) general organization of live animal; (B) 


detail of the cephalic area; (C, D) detail of post-pharyngeal area: based on holotype (C), and paratype (D); (EB) 
copulatory stylet; (F) accessory stylet (both from holotype); (G) idiogram. See Fig. 1 for explanation of abbre- 


viations. 


seminal vesicles, connected to a copulatory 
bulb, is the plesiomorphic condition for the 
Unguiphora (Curini-Galletti & Martens 
1992). The condition seen in A. mirabilis 
may thus have involved a separation of the 
seminal vesicles and their connection to a 
copulatory stylet. The presence of two cop- 
ulatory organs plausibly justifies the exis- 
tence of the large male antrum, similarly 
unknown in other Unguiphora. The extreme 
reduction of the copulatory bulb is a further 
character peculiar to A. mirabilis. 

In addition, among Unguiphora, acces- 


sory stylets are only known in the genus 
Polystyliphora, where, however, they are 
serially arranged, up to 55 in number, mor- 
phologically distinct from the copulatory 
stylet, and connected to a glandular (“‘pros- 
tatoid’’) organ (Curini-Galletti 1998, Curi- 
ni-Galletti & Martens 1991). In A. mirabi- 
lis, copulatory and accessory stylets are 
morphologically identical, and the few ac- 
cessory stylets present do not appear to be 
connected to any glandular structure. Their 
function is puzzling. They may act as a res- 
ervoir of functional stylets and replace the 


VOLUME 114, NUMBER 3 


Fig. 4. 


(A) Nematoplana indica, new species, stylet (holotype). (B) Alloeostyliphora mirabilis, new genus, 


new species, post-pharyngeal area (holotype: lactophenol whole mount, contracted after fixation). (C, D) Ne- 
matoplana calamus, new species, stylet: (C) from holotype; (D) from paratype. 


copulatory stylet once it is damaged or dis- 
charged. Should this be the case, the holo- 
type, which possesses one of the copulatory 
organs with two copulatory stylets con- 
nected to the same seminal vesicle, may 
thus show the transitory stage of the con- 
nection of the first accessory stylet to the 
seminal vesicle, before the former copula- 
tory stylet is discharged. 

The eye-spots of Alloeostyliphora mira- 


bilis appear identical to those found in a 
few species of Nematoplana (Curini-Gal- 
letti & Martens 1992). Their occurrence 
supports the hypothesis of the plesiomorph- 
ic nature of the character in the Unguipho- 
ra, proposed by Ehlers & Sopott-Ehlers 
1990. 

The phylogenetic relationships of Al- 
loeostyliphora mirabilis are unclear. Pro- 
posal of a sister-group relationship with the 


744 


genus Polystyliphora would imply the ho- 
mology of the ‘prostatoid stylets’ with the 
‘accessory stylets’ of A. mirabilis, which, 
at the moment, does not appear to be sup- 
ported. In fact, the morphology of the ‘ac- 
cessory stylets’ (claw-shaped, without 
apophysis) appears uniquely derived for the 
Unguiphora and is not phylogenetically in- 
formative. However, it is parsimonious to 
assume that the additional stylets (with a 
shifting of function from ‘replacement’ (?) 
to ‘prostatoid’, or vice versa) may have 
evolved only once in the Unguiphora. Find- 
ings of additional species of the group may 
shed light on the question. 


General Considerations 


Although accuracy of the morphological 
descriptions was in some cases hampered 
by the small number of specimens available 
for study, the sample was, nonetheless, 
worthy of interest due to the paucity of data 
from extra-European areas and for its spe- 
cific and phylogenetic diversity. The pres- 
ence of three unguiphorid species in a very 
small sample of sediment is surprising and 
points to a remarkable local diversity of 
proseriate fauna. For comparisons, the 
whole of northern Europe harbors only two 
unguiphorid species, while three species are 
known for the Mediterranean basin, where, 
however, each species occupies a distinct 
habitat and never co-occurs in the same 
sediment (Curini-Galletti & Martens 1991, 
1992). 

The finding of species showing unsus- 
pected features, and whose phylogenetic re- 
lationships can not be assessed at present, 
clearly reflects the inadequate state of our 
knowledge of proseriate diversity, especial- 
ly from tropical areas. A peculiar derived 
feature, which is apparently widespread in 
indo-pacific unguiphorids, but not else- 
where, is the presence of unpaired (in cases 
only one) oocytes in front of the pharynx 
(Curini-Galletti et al. 2001, present paper). 
There is no evidence of any further shared 
synapomorphyies among these species, and 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the character does not appear to have a phy- 
logenetic base. A common adaptive strate- 
gy may, however, be involved. In constrast 
to the prevalently stocky and torpid, boreal 
unguiphorids, most tropical unguiphorids 
appear markedly smaller and more slender. 
Differential timing of maturity of oocytes 
on the two sides of the body might be due 
to morphological constraints of the egg size 
in tiny, exceedingly filiform, organisms. 


Acknowledgments 


We are grateful to Donatella Ciaccafava 
Curini-Galletti, who collected and trans- 
ported the sediment sample. 


Literature Cited 


Ax, P, 1958. Vervielfachung des mannlichen Kopu- 
lationsapparates bei Turbellarien.— Verhandlun- 
gen der Deutschen Zoologischen Gesellschaft in 
Graz, pp. 227-249. 

Curini-Galletti, M. 1998. The genus Polystyliphora 
Ax, 1958 (Platyhelminthes: Proseriata) in east- 
ern Australia—Journal of Natural History 32: 
473-499. 

, & P. M. Martens. 1991. Systematics of the 

Unguiphora (Platyhelminthes: Proseriata) I. Ge- 

nus Polystyliphora Ax, 1958.—Journal of Nat- 

ural History 25:1089—1100. 

, & . 1992. Systematics of the Ungui- 

phora (Platyhelminthes: Proseriata) II. Family 

Nematoplanidae Meixner, 1938.—Journal of 

Natural History 26:285—302. 

, G. Oggiano, & M. Casu. 2001. The genus 

Nematoplana Meixner, 1938 (Platyhelminthes: 

Unguiphora) in eastern Australia—Journal of 

Natural History (in press). 

, I. Puccinelli, & P M. Martens. 1989. Kary- 
ometrical analysis of 10 species of the subfam- 
ily Monocelidinae (Proseriata, Platyhelmin- 
thes) with remarks on the karyological evolu- 
tion of the Monocelididae.—Genetica 78:169— 
178. 

Ehlers, U., & B. Sopott-Ehlers. 1990. Organization of 
Statocysts in the Otoplanidae (Plathelminthes): 
an ultrastructural analysis with implications for 
the phylogeny of the Proseriata.—Zoomorphol- 
ogy 109:309-318. 

Marcus, E., 1949. Turbellaria Brasileiros (7).—Bole- 
tins da Faculdade de Filosofia, Ciencias e Le- 
tras, Universidade de S. Paulo, Zoologia, 14:7— 
156. 

Martens, P. M. 1984. Comparisons of three different 


VOLUME 114, NUMBER 3 TAS 


extraction methods for Turbellaria.—Marine gemeiner Teil).—Tierwelt Nordund Ostsee 33, 
Ecology Progress Series 14:229—283. Teil [Vb: 1-146. 

, M. Curini-Galletti, & I. Puccinelli. 1989, On  ‘Tajika, K. I., 1979. Marine Turbellarien aus Hokkaido, 
the morphology and karyology of the genus Ar- Japan III. Nematoplana Meixner, 1938 (Proser- 
chilopsis (Meixner) (Platyhelminthes, Proseria- iata, Nematoplanidae).—Journal of the Faculty 
ta)—Hydrobiologia 175:237—256. of Science of the Hokkaido University, Series 


Meixner, J. 1938. Turbellaria (Strudelwiirmer), I (AlIl- VI, Zoology, 22:69—87. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):746—787. 2001. 


Studies on western Atlantic Octocorallia (Coelenterata: Anthozoa). 
Part 1: The genus Chrysogorgia Duchassaing & Michelotti, 1864 


Stephen D. Cairns 


Department of Systematic Biology—Invertebrate Zoology, National Museum of Natural History, 


Smithsonian Institution, Washington, D. C. 20560-0163, U.S.A. 


Abstract.—The nine species of Chrysogorgia known from the western At- 
lantic are described and illustrated. One species is described as new (C. her- 
dendorfi), three species are synonymized (C. orientalis, C. affinis, C. elisa- 
bethae), and two species are elevated in rank from variety to species (C. 
multiflora and C. thyrsiformis). Lectotypes are chosen for six of the species 
and a neotype for one, C. desbonni, the type species of the genus. A type 
species is also designated for the genus Dasygorgia, a junior synonym of 
Chrysogorgia. An identification key to the nine species is given, as well as 
a list of the 59 currently recognized species in the genus, along with their 
type locality, depth of capture, and branching sequence. The study was based 
on newly reported specimens from over 150 deep-water stations as well as 
re-examination of all historical material; additional records of C. squamata 
and C. spiculosa represent the first reports of these species since their original 


description in 1883. 


Species of the genus Chrysogorgia are 
reputed to be among the most interesting 
and beautiful of the gorgonians (Agassiz 
1888; Verrill 1883), their golden, iridescent 
branches and mathematically precise 
branching pattern making them truly ele- 
gant organisms. They occur exclusively in 
deep water (100-3375 m) and are found 
worldwide except off Antarctica. Members 
of this genus are extremely diverse in col- 
ony form and sclerite complement, the 59 
recognized species (Table 1) representing 
the most speciose of the calcaxonian gen- 
era. The western Atlantic Chrysogorgia 
were reviewed by Verrill (1883) and Deich- 
mann (1936); however, significant collec- 
tions made after 1936 and improved meth- 
ods of study have made a faunistic revision 
of this genus desirable. Indeed, the discov- 
ery of one such undescribed “‘golden cor- 
als” from the gold-carrying wreck of the 
S.S. Central America was the original mo- 
tivation for this paper. 


Materials and Methods 


This study was based on an examination 
of previously unreported specimens collect- 
ed from 157 stations within the western At- 
lantic (Table 2), most collected after Deich- 
mann’s (1936) classic revision of the west- 
ern Atlantic Octocorallia. These specimens 
are deposited primarily at the USNM and 
MCZ, the USNM material originating pri- 
marily from the prodigious collecting of the 
Rosenstiel School of Marine and Atmo- 
spheric Science, University of Miami in the 
1970’s. Historically important specimens, 
including the types for all species dis- 
cussed, were also examined; these speci- 
mens are deposited at the MCZ, USNM, 
and BM. 

The terminology used for the descrip- 
tions follows Bayer, Grasshoff & Versev- 
eldt (1983), whereas general methodology 
of specimen examination can be found in 
Bayer (1961) and Alderslade (1998). Syn- 
onymies for all species are purported to be 


VOLUME 114, NUMBER 3 


Table 1.—The 59 valid species of the genus Chrysogorgia, arranged by group, and within group by branching 
sequence and then date of description. Species from Western Atlantic in bold face. Type localities and depths 


also 


included. 


Group A: “‘Spiculosae’’ (rods and/or spindles in body wall and tentacles) 


C- 
C. 
C. 
C. 
C. 
C. 


OVO ore) © 


C. 
C. flexilis var. maldivensis 


LAO Ore 


C. 


SEOUS ESOL One 


cupressa (Wright and Studer, 1889) 
lata Versluys, 1902 

terasticha Versluys, 1902 
pusilla Versluys, 1902 

dispersa Kiikenthal, 1908 
pyramidalis Kiikenthal, 1908 
=C. aurea Kinoshita, 1913 
rotunda Kinoshita, 1913 
papillosa Kinoshita, 1913 
minuta Kinoshita, 1913 
okinosensis Kinoshita, 1913 
comans Kinoshita, 1913 
sphaerica Aurivillius, 1931 
desbonni Duch. & Mich., 1864 
=C. occidentalis Versluys, 1902 
spiculosa (Verrill, 1883) 
elegans (Verrill, 1883) 

=C. affinis Versluys, 1902 
fewkesii Verrill, 1883 

flexilis typica 

(Wright & Studer, 1889) 

flexilis var. africana Kiikenthal, 1908 


Hickson, 1940 

pentasticha Versluys, 1902 
anastomosans Versluys, 1902 
arborescens Nutting, 1908 

debilis Kiikenthal, 1908 

quadruplex Thomson, 1927 
multiflora Deichmann, 1936 
thyrsiformis Deichmann, 1936 

=C. elisabethae Bayer, 1951 
herdendorfi, n. sp 

pellucida Kikenthal, 1908 
squarrosa (Wright. & Studer, 1889) 
orientalis Versluys, 1902 

mixta Versluys, 1902 

indica Thomson & Henderson, 1906 
irregularis 

Thomson & Henderson, 1906 
dichotoma 

Thomson & Henderson, 1906 


(Banda Sea, 256 m) 
(Celebes Sea, 1901 m) 
(Banda Sea, 204 m) 
(Timor Sea, 520 m) 
(Japan, depth?) 

(Japan, ““upper abyssal’’) 


(Japan, 732 m) 

(Japan, 730 m) 

(Japan, 146 m) 

(Japan, 366 m) 

(Japan, 146 m) 

(Japan, 183-732 m) 
(Lesser Antilles, 161 m) 


(Lesser Antilles, 991 m) 
(Barbados, 433 m) 


(Lesser Antilles, 1048 m) 
(Southern Chile, 220 m) 


(East Africa, ““upper abyssal’’) 


(Maldive Ids., 229-914 m) 


(Timor Sea, 520 m) 
(Indonesia, 520—827 m) 
(Hawaii, 722—914 m) 
(Japan, depth?) 
(Azores, 1022 m) 
(Lesser Antilles, 991 m) 
(Barbados, 183 m) 


(off S. Carolina, 2178 m) 
(Japan, 100 m) 
(Philippines, 914 m) 
(Indonesia, 918—924 m) 
(Celebes, 1165-1264 m) 
(Sri Lanka, 1038 m) 

(Sri Lanka, 1271 m) 


(Bay of Bengal, 165 m) 


747 


1/4L 
1/4L 
1/4L 
1/4L 
1/4L 
1/4L 


1/4L 
1/4L 
1/4L 
1/4L 
1/4L 
1/4L 
2/5R (biflabellate) 


2/5R 
2/5R 


2/5R 
2/5R 


2/SR 
2/5R 


2/5R 
2/5R 
2/5R 
2/5R 
2/5R 
2/5R 
2/5R 


2/5R-3/8R 
dichotomous 
lreg. 

reg. 

lreg. 

reg. 

ireg. (1/3) 


irreg. (L) 


Group B: “‘Squamosae aberrantes’’ (rods and/or spindles in tentacles but not in body wall) 


E 
. expansa (Wright & Studer, 1889) 
. octogonos Versluys, 1902 

. versluysi Kinoshita, 1913 

. calypso Bayer & Stefani, 1988 

. intermedia Versluys, 1902 

. Squamata (Verrill, 1883) 

. bracteata Bayer & Stefani, 1988 
. admete Bayer & Stefani, 1988 


GES) 


AAA A © 


flavescens Nutting, 1908 


(Hawaii, 1688-1977 m) 
(Kermadec Ids., 951 m) 
(Timor Sea, 520 m) 
(Japan, 732 m) 
(Celebes, 732 m) 
(Timor Sea, 250 m) 
(Lesser Antilles, 431 m) 
(Philippines, 329 m) 
(New Caledonia, 390 m) 


1/3L 
1/4R 
1/4R 
1/4R 
1/4R 
1/4R-—1/7R 
1/5R-1/7R 
1/6R 
1/7R 


7A8 
Table 1.—Continued. 


C. agassizii (Verrill, 1883) 

C. curvata Versluys, 1902 

C. chryseis Bayer & Stefani, 1988 
C. stellata Nutting, 1908 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Group C: “‘Squamosae typicae’’ (rods and/or spindles not present; only scales) 


C. axillaris (Wright & Studer, 1889) 
C. geniculata (Wright & Studer, 1889) 
C. fruticosa (Studer, 1894) 

rigida Versluys, 1902 

sibogae Versluys, 1902 

excavata Kiikenthal, 1908 

delicata Nutting, 1908 

cavea Kinoshita, 1913 

ramosa Versluys, 1902 

acanthella (Wright & Studer, 1889) 
pendula Versluys, 1902 

campanula Madsen, 1944 

electra Bayer & Stefani, 1988 
scintillans Bayer & Stefani, 1988 
C. japonica (Wright & Studer, 1889) 


WOO ONDA AOD © 


complete, including every reference to the 
species discussed. In the material examined 
sections for each species, the station num- 
ber is followed by the number of specimens 
in that lot and then the catalog number. The 
SEM photomicrographs were taken by the 
author using an AMRAY 1810 scanning 
electron microscope. 

The following abbreviations are used: 
Vessels: Alb—U. S. Fish Commission 
Steamer Albatross; Atl—Atlantis and R/V 
Atlantis IT; BL—U. S. Coast Survey Steam- 
er Blake; CI—R/V Columbus Iselin; G—R/ 
V Gerda; Gos—R/V Gosnold; GS—R/V 
Gilliss; O—M/V, R/V Oregon and R/V 
Oregon IT; P—R/V Pillsbury; SB—M/V, R/ 
V Silver Bay. 

Museums: BM—The Natural History 
Museum (London); MBD BR—Blake 
Ridge Expedition of the Museum of Bio- 
logical Diversity (The Ohio State Univer- 
sity, Columbus); MCZ—Museum of Com- 
parative Zoology, Harvard (Cambridge); 
USNM—United States National Museum 
(now known as the National Museum of 
Natural History), Washington D. C. 


(Georges Bank., 2271 m) 2/5R 
(Indonesia, 1089 m) biflabellate 
(Ceram Sea, 732 m) biflabellate 
(Hawaii, 649-678 m) multiflabellate 
(Phil./Kerm., 150, 1097 m) 1/3L 
(Philippines, 150-187 m) 1/3L 
(Gulf of Panama, 837 m) 1/3L 
(Philippines, 522 m) 1/3L 
(Banda Sea, 204 m) 1/3L 
(Japan, “‘abyssal’’) 1/3L 
(Hawaii, 536-1463 m) 1/3L 
(Japan, 640-752 m) 1/3L 
(Philippines, 522 m) 1/4L 
(Kermadec, 1097 m) 2/5L 
(Banda Sea, 1595 m) 2/5 
(Iceland, 2448 m) 2/5L 
(Paumotu Ids., 1485 m) flabellate 
(Hawaii, 1758-1937 m) flabellate 
(Japan, 3375 m) unknown 


Subclass Octocorallia 
Order Alcyonacea 
Suborder Calcaxonia Grasshoff, 1999 
Family Chrysogorgiidae Verrill, 1883 


Chrysogorgidae Verrill, 1883: 21.—Nut- 
ting, 1908: 587. 

Chrysogorgiidae Versluys, 1902: 2—4.— 
Kiikenthal, 1919: 486—496 (key to gen- 
era); 1924: 38 (key to genera).—Deich- 
mann, 1936: 220—222 (key to genera).— 
Hickson, 1940: 306—307.—Madsen, 
1944: 44.—Bayer, 1949: 237-238; 1956: 
F216; 1961: 297-299 (key to genera); 
1973: 84-86; 1974: 262—263.—Bayer & 
Muzik, 1976: 67—69 (key to genera).— 
Bayer, 1979: 876-878 (key to genera).— 
Bayer & Stefani, 1988: 257—259 (key to 
genera).—Williams, 1992a: 379; 1992b: 
Die 

Dasygorgidae Studer, 1887: 39.—Wright & 
Studer, 1889: xxxix, 1. 

Dasygorgiidae Hickson, 1904: 221.— 
Thomson & Henderson, 1906: 26. 

Malacogorgiidae Hickson, 1904: 226. 


Diagnosis.—Calcaxonians having an un- 
jointed, solid (non-spicular), concentrically 


749 


VOLUME 114, NUMBER 3 


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754 


Table 2.—Continued. 


Date 


Depth (m) 


Longitude (°’”W) 


Latitude (°’”N) 


Station 


15 VII 1970 
17 VII 1971 


805-1089 


180 


77 34 48 
68 52 54 


17 21 24 
20 11 00 
MV, R/V Silver Bay (SB) 


1262 
1410 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


8 VI 1958 
4 XI 1961 


439-503 
265 


TS) 5) 


27 21 N 
20 53 


440 
3497 


73 44 


layered scleroproteinous axis. The axis of 
stem and branches may be branched or un- 
branched, arising from a root-like or discoi- 
dal, strongly calcified holdfast. The axial lay- 
ers are not undulated and the axial surface is 
smooth (not longitudinally grooved); the axis 
usually displays metallic or iridescent reflec- 
tions. Polyps contractile but not retractile, ar- 
ranged in rows, never in whorls or on op- 
posite sides of branch. Sclerites predomi- 
nantly flat, smooth scales and plates, in some 
species accompanied by warty rods or spin- 
dles. Scales show concentric bands of inter- 
ference colors in polarized light. 

Remarks.—The Chrysogorgiidae is one of 
five families placed in the newly created sub- 
order Calcaxonia (Grasshoff 1999). Previous- 
ly placed in the suborder Holaxonia, these 
families differ in having an axis that does not 
have a cross-chambered, hollow central core, 
but rather a solid axis containing abundant 
calcareous material that is embedded in gor- 
gonin or as purely calcareous segments. The 
taxonomic history of the family is integrated 
into the generic account. 


Subfamily Chrysogorgiinae Verrill, 1883 


Chrysogorginae Studer, 1887: 41.—Wright 
& Studer, 1889: xl, 4. 

Chrysogorgiinae Versluys, 1902: 4, 17.— 
Nutting, 1908: 588.—Madsen, 1944: 
49.—Bayer, 1956: F216. 


Diagnosis.—Chrysogorgiids having 
branching colonies. 

Remarks.—Two other subfamilies are cur- 
rently recognized: the Lepidogorgiidae, for 
species having unbranched colonies and un- 
iserially arranged polyps, and the Chalcogor- 
giinae, for species having unbranched colo- 
nies and polyps arranged bi- or multiserially. 
However, over the last several decades, su- 
bfamilial classification has rarely been used. 


Genus Chrysogorgia Duchassaing & 
Michelotti, 1864 


Chrysogorgia Duchassaing & Michelotti, 
1864: 13 (107).—Verrill, 1883: 21.—Stu- 


VOLUME 114, NUMBER 3 


der, 1887: 41.—Wright & Studer, 1889: xli, 
23.—Versluys, 1902: 17—33.—Nutting, 
1908: 588.—\Ktikenthal, 1919: 505-511 
(key to species); 1924: 388-390 (key to 
species).—Deichmann, 1936: 227-228 
(key to w. Atlantic species)—Hickson, 
1940: 307.—Madsen, 1944: 49.—Bayer, 
1956: F216.—Bayer & Stefani, 1988: 259 
(key to species of ““Squamosae aberran- 
tes””).—Williams, 1992b: 252. 
Dasygorgia Verrill, 1883: 21.—Studer,. 1887: 
41.—Wright & Studer, 1889: xli, 6-9, 278. 


Diagnosis.—Sympodially branched 
chrysogorgiids, the branches. subdividing 
dichotomously, either arising from a regular 
single, ascending spiral around the main 
stem or forming two parallel fans above a 
short main stem. Polyps large relative to 
branches on which they sit, few in number, 
and well separated from one another. Scler- 
ites consist of spindles, rods and scales. 
Axis with a brilliant metallic luster, usually 
amber, yellow or golden in color. 

Discussion.—Verrill (1883) established 
the family Chrysogorgiidae for three gen- 
era: Chrysogorgia, Dasygorgia, and Irido- 
gorgia. He distinguished Dasygorgia from 
Chrysogorgia mainly by its longitudinal ar- 
rangement of sclerites in the body wall, 
Chrysogorgia having transversely and often 
curved sclerites. He included five newly de- 
scribed species in Dasygorgia but did not 
designate a types species. Thus, the first 
species described by Verrill, D. agassizii, is 
herein designated the type of that genus. In 
that paper Verrill (1883) described a major- 
ity of the chrysogorgiid species now known 
from the western Atlantic. Shortly thereaf- 
ter, Studer (1887) unnecessarily rechris- 
tened the family Dasygorgidae, arguing that 
the type species of Chrysogorgia, C. des- 
bonni, could not be correctly identified. He 
also divided the family into two subfami- 
lies, the Chrysogorginae containing all gen- 
era having branched colonies and thus in- 
cluding Chrysogorgia. Two years later, 
Wright & Studer (1889) informally sug- 
gested two groupings among the species of 


55 


Dasygorgia: the “‘Spiculosae’”’ (species 
having spindles and/or rods) and the 
*“Squamosae”’ (species having predomi- 
nantly scale-like sclerites). 

Based on the chrysogorgiids collected on 
the Siboga expedition, Versluys (1902) essen- 
tially monographed the genus Chrysogorgia, 
describing or redescribing the 36 species 
known at that time and presenting a lengthy 
and thorough discussion of the morphology 
of the species in this genus. He convincingly 
demonstrated that Dasygorgia was a junior 
synonym of Chrysogorgia, making Dasygor- 
giudae subordinate to Chrysogorgiidae. He 
adopted the species groupings suggested by 
Wright & Studer (1889), added a third group 
with intermediate characteristics, and formal- 
ized the diagnoses on these groupings. Thus, 
the “Spiculosae”’ (also called Group A) con- 
tained species having rods and/or spindles in 
both the body wall and tentacles, the ““Squa- 
mosae typicae” (also called Group C) con- 
tained species having sclerites in the form of 
scales in both body wall and tentacles, and 
Versluys’ new grouping, the “Squamosae 
aberrantes”’ (also called Group B), contained 
species having scales in the body wall, but 
rods and/or spindles in the tentacles. Versluys 
further subdivided each of these groupings 
into two to four subgroups based on the 
branching sequence of each species. AI- 
though Kiikenthal (1919, 1924) used the 
characteristics of these subgroupings in his 
species key, no author has ever suggested that 
the three major groupings or nine subgroup- 
ings be used as the basis for subgeneric taxa. 
Other significant works concerning the tax- 
onomy of the species of Chrysogorgia in- 
clude: Kinoshita (1913), a review of the Jap- 
anese species; Deichmann (1936), a review 
of the western Atlantic species; Madsen 
(1944), the Ingolf specimens from off Ice- 
land; Bayer (1973), ecological remarks; and 
Bayer & Stefani (1988), new species from the 
western Pacific. 

Type species.—Chrysogorgia desbonni 
Duchassaing & Michelotti, 1864, by mon- 
otypy. 


756 


Fig. 1. Chrysogorgia herdendorfi, holotype, 
USNM 91934, height 20 cm. 


Type species of Dasygorgia: D. agassizii 
Verrill, 1883, here designated. 


Chrysogorgia herdendorfi, new species 
Figs. 1-2 


Description.—The holotype is 20 cm in 
height and about 3 cm in width, the colony 
being of a bottlebrush shape. Colonies are 
attached to the substrate by a roughly cir- 
cular, thin, encrusting (not rhizoidal) hold- 
fast 5-10 mm in diameter, the holdfast be- 
ing milky white in color. The main stem 
just above the holdfast is circular in cross 
section, 0.9—1.1 mm in diameter, and of a 
metallic golden-brown luster. The main 
stem ascends in a tight, counterclockwise 
spiral (producing a zig-zag pattern), giving 
off branches in a regular manner in a pre- 
dominantly 2/5R sequence; however, in 
parts of the colony a 3/8R sequence appears 
to hold. The lowest branch of the holotype 
occurs 16 mm above the holdfast, whereas 
the lowest branch of the paratype is at 60 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


mm height, but bears scars of broken 
branches as low as 30 mm. Branches occur 
approximately every 2.5—3.0 mm along the 
main stem, the distance between aligned 
branches (defined as the “‘orthostiche inter- 
val” by Versluys 1902) in the 2/5R se- 
quence being 14-17 mm. The lowermost 
branches are often unbranched, up to 32 
mm in length, and bear up to 6 polyps. The 
branches of the mid-portion of the colony 
usually have only one node, the first inter- 
node 6—7 mm in length, the terminal twigs 
up to 35 mm, producing a total branch 
length of about 42 mm. One polyp occurs 
on the first internode at or within 0.5 mm 
of the node, and 4—6 polyps occur on the 
terminal twigs. Branches toward the top of 
the colony usually have two nodes, the first 
internode about 4 mm in length, the second 
6—7 mm, and the terminal twigs up to 30 
mm, also producing a total branch length of 
about 42 mm. There is usually one polyp 
on the first and second internodes adjacent 
to the nodes, and 4—5 polyps on the ter- 
minal twigs. Branches near the growing tip 
of the colony also usually have two nodes, 
but the terminal twigs are shorter (about 10 
mm), thus result in shorter branches. Hence, 
most branches of the colony are of the same 
total length, i.e., 40-45 mm, regardless of 
the number of internodes present or the lo- 
cation of the branch, except for those near 
the developing tip, which are shorter. No 
branches containing three internodes were 
noted. The branches arise at gradually in- 
creasing angles from the base upward, the 
lowest branches about 45° from the vertical, 
those toward the top of the colony 60—70° 
from the vertical. Branch internodes adja- 
cent to the main stem are quite slender, 
about 0.08 mm in diameter; distal twig 
branch diameters are even smaller, about 
0.04 mm. These slender branches produce 
a very flexible limp structure to the colony, 
such that when a colony is removed from 
fluid the branches will coalesce with one 
another. Corresponding internodes and ter- 
minal twigs of the same branch are of the 
same lengths. There is no anastomosis. 


USNM 91934. A, six spindles from body wall 


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Fig. 2. 
and tentacles; B, 


VOLUME 114, NUMBER 3 


758 


The polyps are elongate, up to 2.3 mm 
in height and 0.6 mm in width, slightly con- 
stricted at mid-level, and diverge at an 
oblique angle to the branch. The thin coe- 
nenchyme covering both the main stem and 
branches contains straight, elongate scales 
that are usually blunt at each end, up to 0.43 
mm in length and 0.06—0.07 mm in width. 
These scales are have finely serrated edges 
and bear very small granules (1.8—2.5 wm 
diameter) on their faces. Toward the base 
of the polyps and continuing up the body 
wall the scales gradually change into lon- 
gitudinally oriented, straight spindles of vir- 
tually the same size; the spindles are point- 
ed at both ends and bear small compound 
warts, the warts 4—8 «zm in diameter. Thus 
the body wall consists of both elongate 
scales and spindles of similar size and 
shape. The lower, backs of the tentacles also 
contain robust, finely-warted spindles, usu- 
ally arranged three across a tentacle, up to 
0.43 mm in length and 0.11 mm in width. 
The pinnules contain small, irregularly-rect- 
angular scales, most about 0.15—0.20 mm 
in length. They have finely-serrated edges 
and virtually flat faces that bear very small 
granules, the largest only | 4m in diameter. 

Discussion.—At first glance, the branch- 
ing sequence of C. herdendorfi appears to 
be a typical 2/5R, like the morphologically 
similar C. elegans. This branching formula, 
a shorthand notation developed by Versluys 
(1902), implies that five branches originate 
from the main stem in two counterclock- 
wise revolutions up the main stem (i.e., 
counterclockwise, as viewed from the apex 
of the colony; and as moving to the 
“right’’, as viewed for the side of the col- 
ony), such that every sixth branch will lie 
directly above the first, forming five rows 
of branches along the main stem. However, 
the branch correspondence was often found 
to be slightly off after two revolutions up 
the stem. But when branch origins were 
traced for three revolutions around the main 
stem, an exact correspondence was discov- 
ered, such that the branch sequence would 
appear to be 3/8R. This implies that there 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


are eight longitudinal rows of branches 
from the main stem, but that three revolu- 
tions of branch origins must occur before a 
branch (the 9th) lies directly above the first. 
The angular separation between adjacent 
branches is thus 135°, instead of 144°, the 
latter being typical of the 2/5 sequence. 
This is believed to be the first example of 
a chrysogorgiid branching sequence that re- 
quires three revolutions to achieve align- 
ment. 

Although two specimens are unquestion- 
ably assigned to this species, both from the 
wreck of the Central America, five Chry- 
sogorgia colonies were collected from that 
site. Two of the other three specimens 
(USNM 91935 and BR127) are identical to 
C. herdendorfi in all characters described 
above, except that their body wall sclerites 
are longitudinally arranged, elongate scales, 
much like those in the lower body wall of 
C. herdendorfi, but never graduating to the 
spindle shape characteristic of the upper 
body wall of C. herdendorfi. This may rep- 
resent variation in sclerite form, or a close- 
ly-related undescribed species, but, because 
of the paucity of specimens available and 
the uncertainty in interpreting this character, 
they are presented here as a non-type vari- 
ety of C. herdendorfi. The fifth specimen 
(BR234) is even more perplexing in that it 
has sclerites similar to C. herdendorfi but it 
is a larger colony (36 cm height) with more 
nodes (0-2-5) and longer, flabellate branch- 
es. It may well represent an undescribed 
species and is probably the species most of- 
ten figured in situ by Herdendorf, et al. 
(1995: 93-94, 111, 189, figs. 62-64, 66— 
67, 71-72, 83, 85; 89, 93-957 990MM Se 0o: 
113 (also back cover), 128-129, 147), fig- 
ure 89 being the actual specimen collected 
(BR234). But, until additional specimens 
are collected and examined, this species 
will remain undescribed. 

Within the western Atlantic, C. herden- 
dorfi is most similar to two species: C. ele- 
gans and C. agassizii (see Table 3). But, 
although similar in colony shape and spic- 
ulation, C. elegans differs in having a more 


759 


VOLUME 114, NUMBER 3 


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760 


Table 3.—Continued. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


petite colony with much more closely 
spaced branches, a rhizoidal holdfast, a 2/ 
5R branching sequence, branches with pre- 
dominantly two nodes, relatively short ter- 
minal twigs, thicker terminal twigs that pro- 
duce a wiry (not limp) colony, and larger 
and more rotund body wall sclerites (rods 


Distribution and depth range 
in Western Atlantic 
off Georges Bank 
1928-3700 m 
Antilles, Yucatan, 
431-1046 m 


ag ae instead of spindles). Also, C. elegans is 
eee Ee Svalieogte known only to the south of C. herdendorfi 
83 Z 4 & g es e : 5 and at lesser depths. C. agassizii is similar 
Eee @ 5 iors 39 in colony shape and branch flexibility, but 
Base Bh g 6 So S differs in having more closely-spaced 
e2e° = = g so branches, longer branches with predomi- 
Epi) PEO EE eo. nantly three nodes, slightly larger tentacular 
235 2 5 2 2 5 = 3 3 5 spindles, and the absence of “true spicules” 
(i.e., spindles or rods) in the body wall. The 

2 San 4 exclusive presence of flattened scales in the 
gue] 5 s body wall of C. agassizii, in fact, places it 
S in the a different subgroup of the genus, the 

BN EB ““Squamosae aberrantes’’. 

O Etymology.—This species is named in 
SEs honor of Charles E. Herdendorf, biological 
ae S S coordinator aboard the R/V Arctic Discov- 
Zoe if erer during its recovery operations of the 

z = ep S.S. Central America. 

eer a Material examined (Types).—Holotype: 

oes USNM 91934, 1 colony, 31°35’ N, 
Eee | “6 TT°10'W, (270 km off Charleston, South 
Bue @ : e : Carolina), 2178 m, August 1990; Paratype: 
532 a a MBD BR-128, 1 colony, locality as above. 
: aa r Type Locality: wreck of the S.S. Central 
Bm) : E : America (collected by the research sub- 
PaaG| oS = oo mersible “‘Nemo” operated by the R/V Arc- 
see qe fea 2 tic Discoverer): 31°35'N, 77°10'W (approx. 
A a N 270 km off coast of South Carolina), 2178 
m. 
5: u ade Distribution.—Known only from the 
zz| < ar type locality. 
om + a Chrysogorgia elegans (Verrill, 1883) 
£2| 3 Z Figs. 3-4 
Be: : 
S3 2 5 = 5 Dasygorgia elegans Verrill, 1883: 23. 
ae = Dasygorgia spiculosa Verrill, 1883: 23—24 
(in part: BL-44, 1 of 2 specimens; BL- 
= s 195, 1 of 2 specimens; BL-205, 1 speci- 
2 5 men). 
Sy > Dasygorgia spiculosa: Wright & Studer, 
S S) 1889: 9-10, pl. 4, fig. 1, pl. 5, fig. 1. 


VOLUME 114, NUMBER 3 


761 


Fig. 3. Chrysogorgia elegans. A, colony from O-4812, USNM 52859, 14 cm in height; B, rhizoidal holdfast 
with attached cirripede, Citation-5505, USNM 89100, height 19 mm; C, lectotype, MCZ 4860, BL-283, 12 cm 
in height; D, three fragments of holotype of C. affinis, BM 89.5.27.5, tallest fragment 33 mm in height; E, line 
drawing of polyp from lectotype (courtesy of EK M. Bayer), polyp 1 mm in width. 


Chrysogorgia affinis Versluys, 1902: 47— 
48, figs. 64—65.—Kitikenthal, 1919: 521; 
1924: 396.—Deichmann, 1936: 221-— 
222.—Bayer, 1959: 29. [new synonym] 

Chrysogorgia elegans: Versluys, 1902: 61.— 
not Nutting, 1908: 590.0—Kiikenthal, 1919: 
533, fig. 234; 1924: 405.—Deichmann, 
1936: 231—232, in part: pl. 23, figs. 53-59: 


BL-44 (in part), BL-195 (in part), BL-205, 
BL-283); not pl. 33, fig. 1 (=C. spiculo- 
sa).—Bayer, 1952: 189; 1954: 280 (listed); 
1956: F216, fig. 155, 3a—c, 158,6; 1958: 
389, 390 (listed).—?Tixier-Durivault & 
d’Hondt, 1974: 1409.—Grygier, 1984: 143 
(Oregon-548).—?Grasshoff, 1986: 25.— 
Grygier, 1990: 667. 


762 


?Chrysogorgia flexilis: Thomson, 1927: 22. 

Chrysogorgia agassizii: Deichmann, 1936: 
in part (pl. 34, figs. 1-2: labeled correctly 
as D. elegans in Verrill’s unpublished 
plates). 

Chrysogorgia sp. Grygier, 1984: 143 (P- 
781). 


Description.—Colony bottlebrush- 
shaped, up to 16 cm in height, and pos- 
sessing a rhizoidal holdfast. Main stem up 
to 1.1 mm in diameter, golden-brown in lus- 
ter. Branching sequence consistently 2/5R: 
branches closely spaced, one every 1.0—1.5 
mm; orthostiche interval 0—7.5 mm apart. 
Distal branch diameter 0.2 mm; branches 
wiry. Number of nodes per branch ranges 
from 1—3, but usually 2, producing 4 rela- 
tively short (4—6 mm) terminal twigs; some 
branches have | or even 3 nodes, but they 
are never without at least 1 node. First in- 
ternode 6-8 mm in length; second, 2—5 
mm, and terminal twigs 4—6 mm, produc- 
ing a total branch length that rarely exceeds 
25 mm. Usually one polyp occurs per in- 
ternode, and 1—3 on the terminal twigs. Pol- 
yps up to 2.3 mm in height and 0.8 mm in 
diameter, the base of the polyps sometimes 
swollen. Coenenchymal sclerites sparsely 
warted, slender scales up to 0.65 mm in 
length and 0.09 mm in width, serrate on the 
edges. Body wall and tentacular sclerites 
longitudinally arranged, rotund rods 0.44— 
0.65 mm in length and up to 0.12 mm in 
width, usually straight but sometimes 
slightly curved. Body wall rods evenly 
warted, each compound wart about 7 ~m in 
diameter and composed: of several smaller 
elements, each about 1.8 wm in diameter. 
Pinnular and distal tentacle sclerites rect- 
angular to medially-constricted scales 0.12— 
0.20 mm in greater length and 0.0—0.05 mm 
in width. Their faces are sparsely orna- 
mented and their edges are finely serrate, 
an. apex occurring about every 4.2 wm. 

Discussion.—Nutting (1908) reported C. 
elegans from three Albatross stations in the 
Hawaiian Islands; however, Ktikenthal 
(1919) and most other authors have doubted 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


this identification based on the circumstan- 
tial evidence of its disparate location. These 
three specimens, all deposited at the 
USNM, were examined and confirmed to 
differ from C. elegans in branching pattern 
and spiculation, the Hawaiian specimens 
having only irregularly-shaped scales in 
their body walls, and thus pertaining to a 
different subgroup of Chrysogorgia, the 
*“*Squamosae aberrantes’’. 

The holotype of C. affinis Versluys, orig- 
inally described as D. spiculosa by Wright 
& Studer (1889), was based on a specimen 
10 cm in length but apparently broken into 
three fragments by the time Versluys de- 
scribed it, the largest fragment 6 cm in 
length. The holotype is now in six frag- 
ments, the longest segment 44 mm in 
length. The holotype was examined and 
found to perfectly match the characteristics 
of C. elegans, and thus it considered to be 
a junior synonym, as implied by Bayer 
(1959). C. affinis has been reported only 
from its type locality. 

Deichmann (1936) considered C. affinis 
(=C. elegans) and C. spiculosa to be syn- 
onymous; however, these species can be 
distinguished (see Discussion of C. spicu- 
losa and Table 3). Versluys (1902) reaf- 
firmed that distinction by renaming Wright 
& Studer’s (1889) D. spiculosa as a new 
species, C. affinis (which is C. elegans), 
distinct from C. spiculosa. 

Chrysogorgia elegans is reputed to occur 
in the eastern Atlantic off the Azores, Cape 
Verde, Bay of Biscay, and off Morocco 
(Thomson 1927 as C. flexilis, Tixier-Duri- 
vault & d’Hondt 1974, Grasshoff 1986) at 
depths of 946-3088 m. None of the speci- 
mens on which these reports were based 
was illustrated or described, nor have I ex- 
amined them. Given the great depths of 
capture for some of the specimens and the 
gross similarity of various Atlantic species, 
these eastern Atlantic records remain to be 
confirmed. 

Material examined.—Alb-2397, 3, 
USNM 49741; Ati-3306, 2, MCZ; G-403, 
1, USNM_ 100885; G-1010, 1, USNM 


VOLUME 114, NUMBER 3 


Ve 
vi 
4 


“y i \ 
>" 

Hi 
wee 
an 


Fig. 4. Chrysogorgia elegans, sclerites from P-904, USNM 52865. A, seven rods from body wall; B, four 
coenenchymal scales; C, five pinnular scales. 


764 


100883; Citation-4502, over 30, USNM 
89090; Citation-4504, 1, USNM 89091; Ci- 
tation-4507, 18, USNM 89092; Citation- 
4508, over 100, USNM 89093; Citation- 
4509, over 50, USNM 89094; Citation- 
4510, over 50, USNM 89095; Citation- 
4511, over 50, USNM 89096; Citation- 
4512, 10, USNM 89097; Citation-4513, 3, 
USNM 89098; Citation-4514, 1, USNM 
89099; Citation-5505, 44, USNM 89100; 
Gyre-2387, over 50, USNM 89089; Gyre- 
2429, 3 dry, USNM 89101; Gyre—S36, 3, 
USNM 100729; O-489, 5, USNM 49944; 
O-548, 3, USNM 50027, 52866; O-549, 8, 
USNM 50024; O-4413, 1, USNM 52860; 
O-4729, 2, USNM 52866; O-4812, 1, 
USNM 52859; P-394, 1, USNM 52864; P- 
478, 4, USNM 52863; P-776, 10, USNM 
100884; P-781, 4, USNM 52908 (reported 
by Grygier, 1984 as Chrysogorgia sp.); P- 
784, 1, USNM 55919; P-847, 1, USNM 
52849; P-850, 1, USNM 52861; P-881, 
over 70, USNM 52858; P-904, 3, USNM 
5286530 2- 919 Se WSINMES2909 Se-9S8 302. 
USNM 52862; specimens reported by 
Deichmann (1936); types of D. elegans and 
C. affinis (see below). 

Types.—Verrill based the species on six 
specimens (syntypes) collected at three 
Blake stations: one from BL-260 (unknown 
MCZ catalog number), two from BL-283 
(MCZ 4860), and three from BL-284 (MCZ 
4859), made off Grenada and Barbados at 
depths of 433-636 m. Deichmann (1936) 
restricted the type locality to BL-283, but 
did not choose a lectotype from among the 
five (not two) specimens from that station. 
A lectotype is therefore designated as one 
of the five syntypes collected at BL-283 
(MCZ 4860), making the type locality con- 
sistent with Deichmann’s statement, which 
is: 13°05’05"N, 59°40'50”W (west coast of 
Barbados), 433 m. The remaining four 
specimens from BL-283 (MCZ 4860a) and 
the specimens from BL-284 (MCZ 4859) 
and BL-260 are considered to be paralec- 
totypes; however, the specimen from BL- 
260 appears to be lost, and only one of the 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


three specimens from BL-284 could be 
found. 

The holotype of C. affinis, now in six 
pieces (see Discussion), is deposited at The 
Natural History Museum, London 
(89.5.27.5) Type locality: Challenger-122: 
9°05'S, 34°50’'W (off Pernambuco, Brazil), 
640 m. 

Distribution.—Disjunct distribution: 
northern Gulf of Mexico from Tamaulipas 
Province, Mexico to off Florida Panhandle; 
Little Bahama Bank; southeastern Carib- 
bean throughout Lesser Antilles to border 
of Colombia and Panama; Pernambuco, 
Brazil; 128-1716 m, although most records 
between 500—800 m.? Eastern Atlantic; 
946-3088 m (see text). 


Chrysogorgia spiculosa (Verrill, 1883) 
Figs. 5—6 


Dasygorgia spiculosa Verrill, 1883 (in part: 
1 of 2 specimens from BL-44, both spec- 
imens from BL-190, 1 of 2 specimens 
from BL-195, not BL-205, ?BL-222, ?BL- 
227), 23-24, pl. 2, fig. 5—not Wright & 
Studer, 1889: 9-10 (=C. affinis/elegans). 

Chrysogorgia sp. Agassiz, 1888: 144, fig. 
456.—Bayer, 1973: fig. 18. 

Chrysogorgia spiculosa: Versluys, 1902: 
53, 60, 61.—not Nutting, 1908: 591.— 
Kiikenthal, 1919: 537, fig. 235; 1924: 
407—406.—Bayer & Macintyre, 2001: 
342 (minerology). 

Chrysogorgia elegans: Deichmann, 1936: 
231-232 (in part: pl. 22, fig. 7, pl. 33, fig 
1: BL-44 (Gn part), BL-190, BL-195 Gn 
part), BL-200, and MCZ 4861). 


Description.—Colony bottlebrush- 
shaped (but with rather elongate branches), 
up to 25 cm in height; rhizoidal holdfast. 
Main stem up to 1.5 mm in diameter; gold- 
en-brown in luster. Branching sequence 
consistently 2/5R; branches well spaced, 
one every 3.5—4.5 mm; orthostiche interval 
20—25 mm; branches diverge from main 
stem at 80—90° from the vertical. Terminal 
twig diameter 0.2 mm, producing a rigid to 


VOLUME 114, NUMBER 3 


wiry colony. Number of nodes per branch 
3-6, but usually 4 on a well-developed 
branch. First internode long (15—20 mm); 
second through fourth internodes, 10—15 
mm; terminal twigs can be much longer, re- 
sulting in a total branch length of 70—110 
mm. Internodes and terminal twigs of a 
branch usually oriented in a _ horizontal 
plane, sometimes bent slightly downward 
distally. Three polyps usually present on 
first internode; 2 or 3 on successive inter- 
nodes; 2—8 on terminal twigs. Polyps large, 
up to 3.0 mm in height, oriented perpendic- 
ular to branch, and often having a swollen 
basal region that encircles the stem. Coe- 
nenchymal sclerites elongate, slender, 
pointed scales up to 0.60 mm in length and 
0.06 mm in width. Their edges are slightly 
serrate and their faces sparsely warted. 
Body wall sclerites primarily rotund rods 
and sometimes spindles (or rods with one 
blunt end and the other pointed), the rods 
straight to slightly bent, and up to 0.96 mm 
in length and 0.18 mm in diameter. Body 
wall sclerites bear small compound warts 
about 18 wm in diameter, each wart con- 
sisting of 10-15 smaller elements 3.6—4.2 
zm in diameter. Tentacular sclerites also 
rods and spindles, but usually smaller, only 
up to 0.54 mm in length. Pinnular sclerites 
elongate scales about 0.20 mm in length, 
having serrate edges. 
Discussion.—Deichmann (1936) synon- 
ymized C. spiculosa with C. elegans, im- 
plying that even Verrill was uncertain about 
their distinction. In her material examined 
for C. elegans, she listed most (but not all) 
of the type series of both C. spiculosa and 
C. elegans, as well as material from two 
other stations (see synonymy). Deich- 
mann’s confusion was justified, in that Ver- 
rill did include two species (C. elegans and 
C. spiculosa) in his type series of C. spi- 
culosa, both species sometimes even occur- 
ring at the same station (see synonymies of 
respective species). Although C. spiculosa 
is similar to C. elegans in branching se- 
quence, flexibility, and geographic distri- 
bution, it differs in several significant char- 


765 


Fig.5. Chrysogorgia spiculosa. A, colony from G- 
1112, USNM 52854, height 15 cm; B, holdfast from 
G-130, USNM 52856, height 18 mm. 


acters as summarized in Table 3. Specimens 
of C. spiculosa are much more robust, hav- 
ing more widely spaced and longer branch- 
es; more nodes per branch; and more polyps 
per node. Furthermore, the body wall of C. 
spiculosa has much larger rods. Versluys 
(1902) also noted the difference between 
these two species by distinguishing his D. 
affinis (=C. elegans) from D. spiculosa. 
These are the first additional records of C. 
spiculosa since its original description. 
Material examined.—Alb-2751, 1, 
USNM 44109; G-128, 3, USNM 52850; G- 
129, 2, USNM 52851; G-130, 14, USNM 
52856; G-368, 4 dry, USNM 100875; G- 
370, 1, USNM 100876; G-371, 1, USNM 
52855; G-372, 1 dry, USNM 100873; G- 
965, 3, USNM 52907; G-1112, 3, USNM 
52854; GS-134, 1, USNM 100877; O-4701, 
1, USNM 100878; O-4811, 1, USNM 
52852; O-10878, 2, USNM 100879; P-391, 
1, USNM 52853; misidentified specimen of 
Nutting (1908), Alb-4151, USNM 25356; 


width of polyps 1.2—1.3 mm 


b) 


five rods from: body wall; B, four 


> 


scales; EK two polyps 


USNM 52856. A 


; pinnular 


Z 
6 
sal 
© 
g 
= 
Y 
< 
3 
ee 
o) 
> 
ie 
ra 
3) 
fe 
Za) 
— 
< 
S 
©) 
} 
— 
ec 
faa] 
aa 
a 
cal 
ae 
fo) 
Ze) 
0 
g 
al 
jaa 
ea 
9) 
fe) 
~ 
ja 


sogorgia spiculosa, sclerites from G-130, 


, unpublished plates). 


Chry. 
coenenchymal scales; C, a tentacular rod; D—E 


(from Verrill 


a ge ae 


a 


Fig. 6. 


VOLUME 114, NUMBER 3 


specimens listed by Deichmann (1936); 
types of D. spiculosa (see below). 

Types.—Verrill (1883) reported nine 
specimens plus some fragments from six 
Blake stations in his original description, all 
of which must be considered to be synty- 
pes: BL-44, 2, MCZ 4855; BL-190, 2, MCZ 
4856 and 4856a; BL-195, 2, MCZ 4857; 
BL-205, 1, MCZ 4859; BL-222, 1; and BL- 
227, 1. However, because Verrill included 
both C. elegans and C. spiculosa in his syn- 
type series (see synonymies of these two 
species), a lectotype is chosen from BL-190 
(MCZ 4856), a specimen 5.5 cm tall that is 
consistent with the more robust species 
originally envisioned and described by Ver- 
rill. The other syntypes thus become para- 
lectotypes; however, the specimens from 
BL-222 and BL-227 could not be found at 
the MCZ in 2001. Type Locality (as re- 
stricted by lectotype): 15°18'12"N, 
61°26'32”W (off western Dominica, Lesser 
Antilles), 991 m. 

Distribution.—Disjunct distribution: nor- 
thern Gulf of Mexico from Tamaulipas 
Province, Mexico to Havana, Cuba; Lesser 
Antilles from Dominica to St. Vincent; off 
Colombia; 914—2265 m. 


Chrysogorgia multiflora Deichmann, 1936 
new rank 
Figs. 7-8 


Chrysogorgia fewkesi var. multiflora Deich- 
mann, 1936: 231, pl. 22, fig. 6, pl. 23, 
figs. 51-52.—Bayer, 1959: 27-29, fig. 
13a. 


Description.—Colonies bottlebrush- 
shaped, but somewhat bushy; colonies up 
to 25 cm in height. Colony attached by an 
encrusting holdfast; main stem up to 2.1 
mm in diameter, golden-brown in luster. 
Branching sequence consistently 2/5R: 
branches well spaced, one every 3.5—4.0 
mm; orthostiche interval 20—22 mm; how- 
ever, some colonies are diminutive (indi- 
cated with an asterisk in the Material ex- 
amined section), having branches separated 
from one another by only about 1 mm and 


767 


an orthostiche interval of 6 mm. Branches 
usually diverge from main stem at 90—110° 
from the vertical, thus projecting perpen- 
dicular to slightly downward from the di- 
rection of growth, and often giving a false 
impression of the top and bottom of the col- 
ony, especially if the holdfast is missing. 
Branches gently curve upward after the first 
several internodes; fusion of branches oc- 
casionally occurs. First internode robust (up 
to 1.3 mm in diameter), whereas terminal 
twigs are quite delicate (0.1 mm in diame- 
ter), but altogether producing a robust, wiry 
colony. Number of nodes per branch up to 
nine, although usually only 5 or 6, the first 
internode being 10—14 mm in length, sub- 
sequent internodes 5—6 mm in length, and 
terminal twigs of variable lengths, resulting 
in branches up to 90 mm in length. Inter- 
nodes and terminal twigs not arranged in a 
plane but as a bush. Two to 4 polyps usu- 
ally present on first internode; | or 2 on all 
successive internodes; and a variable num- 
ber on the terminal twig, depending on 
length. Polyps small, up to 1.55 mm in 
height. Coenenchymal sclerites of main 
stem elongate, flattened plates, up to 0.65 
mm in length and 0.06 mm in width, bear- 
ing prominent, compound warts up to 24 m 
in diameter. Coenenchymal sclerites of 
branches also flattened plates, but usually 
more elongate (up to 0.77 mm), with point- 
ed ends and less prominent warts. Lower 
body wall sclerites obliquely arranged; up- 
per body wall sclerites transversely ar- 
ranged. Most body wall sclerites curved 
spindles, the curvature corresponding to the 
circumference of the polyp wall, the largest 
spindles up to 1.08 mm in length and 0.07— 
0.13 mm in width, although smaller spin- 
dles are interspersed. Body wall spindles 
sparsely covered with compound warts up 
to 9 wm in diameter. Tentacular sclerites 
longitudinally arranged, curved spindles, 
but usually shorter (up to 0.86 mm). Pin- 
nular sclerites finely granular, often medi- 
ally-constricted, rectangular scales up to 
0.20 mm in length. 
Discussion.—Grasshoff (1981) legiti- 


768 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 7. Chrysogorgia multiflora, colony from G- 
859, USNM 52846, height 20 cm. 


mized and elevated the distinction of vari- 
ety multiflora by synonymizing it with C. 
quadruplex Thompson, 1927, a species 
heretofore known only from the eastern At- 
lantic (Azores, Bay of Biscay, Atlantis Sea- 
mount, Celtic Sea; 507—2682 m) Examina- 
tion of the types of C. fewkesii and variety 
multiflora show small but consistent differ- 
ences (see Discussion of C. fewkesii). The 
synonymy of C. multiflora with C. quad- 
ruplex may be correct but I have not veri- 
fied it. Until this is proven, I choose to rec- 
ognize Deichmann’s western Atlantic vari- 
ety multiflora as a distinct species, but only 
marginally different from C. fewkesii (see 
*“‘Discussion”’ of C. fewkesii and Table 3 for 
their distictions). The records reported here- 
in are the first from the western Atlantic 
since its original description. 

Material examined.—*Alb-2415, 3, 
USNM 44128 and 52841; *Azl-266-40, 1, 
USNM 100880; C/-15, 1, USNM 100881; 
CI-46, 1, USNM 100882; G-130, 4, USNM 
52847; *G-403, 1, USNM 52839; G-859, 
2, USNM 52846; G-1111, 3, USNM 52842; 


*Gos-2385, 1, USNM 56896; O-2081, 4, 
USNM 50907 (reported by Bayer, 1959); 
*O-2636, 1, USNM 51579; P-892, 2, 
USNM 52844; Waldo Schmitt station 65- 
32, off Dry Tortugas, 1064 m, USNM 
50094; specimens reported by Bayer 
(1959); types of C. fewkesii multiflora (see 
below). 

Types.—Deichmann alluded to “some” 
specimens of C. fewkesi var. multiflora from 
Blake-190, which are considered as synty- 
pes (MCZ 4854). These specimens consist 
of several small, poorly-preserved branch- 
es, not including any complete colonies. 
Type locality: 15°18'12"N, 61°26'32’"W (off 
southwestern Dominica, Lesser Antilles), 
991 m. 

Distribution.—Disjunct distribution: off 
mouth of Amazon River, Brazil; Lesser An- 
tilles (St. Christopher to St. Lucia), Straits 
of Florida, and Tongue of the Ocean, Ba- 
hamas; 320—1280 m. 


Chrysogorgia fewkesii Verrill, 1883 
Figs. 9-10 


Chrysogorgia Desbonni: Pourtalés, 1868: 
131-132. 

Chrysogorgia Fewkesii Verrill, 1883: 26. 

Chrysogorgia fewkesii: Wright & Studer, 
1889: 24. 

Chrysogorgia fewkesi: Versluys, 1902: 55— 
56.—Kikenthal, 1919: 533-534; 1924: 
405.—?Thomson, 1927: 21-22, pl. 1, 
figs. 6-~7.—Deichmann, 1936: 222, 230— 
231, pl. 23, figs. 41-50.—Tixier-Duri- 
vault & d’Hondt, 1974: 1409. 


Description.—Colonies bottlebrush- 
shaped, but bushy; colonies up to 22.5 cm 
in height. Nature of holdfast unknown; bas- 
al branch diameter up to 2.9 mm. Branching 
sequence consistently 2/5R; branches well 
spaced, one every 3—6 mm; orthostiche in- 
terval 17-20 mm. Branches diverge from 
main stem at 100—110° from the vertical, 
thus projecting slightly downward at first. 
First internode robust; terminal twigs quite 
narrow, altogether producing a wiry colony. 
Number of nodes per branch usually 5—7, 


VOLUME 114, NUMBER 3 


Fig. 8. Chrysogorgia multiflora. A—D, sclerites from G-859, USNM 52846. A, six curved spindles from 
body wall; B, coenenchymal platelets from side branch; C, coenenchymal platelets from main stem; D, pinnular 
scales. E, two polyps, MCZ “5796”, width of polyps 0.68 mm (from Verrill, unpublished plates). 


770 


Fig. 9. 
lectotype, MCZ 4850, polyps 0.5—0.7 mm in width 
(from Verrill, unpublished plates). 


Chrysogorgia fewkesii, two polyps from 


although may be up to 10, the first inter- 
node being 10—12 mm in length and re- 
maining ones 6—8 mm, resulting in branch 
lengths of up to 100 mm. Internodes and 
terminal twigs not arranged in a plane. Two 
to four polyps occur on each internode, a 
variable number on terminal twigs. Polyps 
small, about | mm in height. Coenenchy- 
mal sclerites of main stem and branches 
flattened, elongate, pointed plates up to 0.95 
mm in length and 0.05—0.06 mm in width, 
having prominent compound warts on their 
faces and edges. Body wall sclerites trans- 
versely arranged, consisting of curved, 
slightly flattened spindles, up to 0.71 mm 
in length and 0.04—0.06 mm in width. Body 
wall spindles bear multiheaded warts up to 
12 wm in diameter and often have one or 
both distal ends strongly flattened. Tentac- 
ular sclerites similar to those of body wall 
but longitudinally arranged and also con- 
taining some shorter blunt rods 0.18—0.30 
mm in length and 0.04—0.06 mm in diam- 
eter. Pinnular sclerites finely granular, rect- 
angular scales 0.10—0.15 mm in greater 
length. 

Discussion.—Chrysogorgia fewkesii and 
C. multiflora, the two western Atlantic spe- 
cies having transverse sclerites in the body 
wall and downward projecting branches, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


are very similar (Table 3). In fact, the col- 
ony form is so similar that the colony figure 
of C. multiflora (Fig. 7) may serve for both 
species. Deichmann (1936) established the 
variety C. fewkesi var. multiflora for several 
specimens collected off Dominica (BL-190) 
that differed from typical C. fewkesii in 
three ways: the variety was purported to 
have a more robust colony, its body wall 
sclerites were “‘better developed’’, and its 
coenenchymal sclerites were larger (0.45 vs 
0.35 mm in length for the typical form) and 
more warty. Concerning point one, there 
seems to be little or no difference in colony 
size between the two species. Concerning 
point two, Deichmann’s illustrations of the 
body wall sclerites of these two taxa 
showed the variety to have slightly wider 
sclerites G.e., 0.07—0.08 mm vs 0.035—0.05 
mm for the typical form) but no difference 
in length. This difference was borne out by 
re-examination of the type material, the 
width of the body wall spindles being even 
greater than Deichmann illustrated G.e., up 
to 0.13 mm). Furthermore, the body wall 
sclerites of C. fewkesii are somewhat short- 
er, slightly flattened, and have very flat- 
tened, rounded distal ends; those of C. mul- 
tiflora are rotund with pointed ends. Con- 
cerning point three, the coenenchymal 
scales of both C. multiflora and C. fewkesii 
are warty and of approximately the same 
size; no difference in degree of ““wartiness”’ 
could be discerned, although it was noted 
that the coenenchymal scales on the main 
stem of C. multiflora were more warty than 
those on the branches. Indeed, the coenen- 
chymal scales of C. fewkesii are, in general, 
longer (not shorter) than those of C. mul- 
tiflora (see Table 3). 

The few specimens of C. fewkesii re- 
ported from the eastern Atlantic by Thom- 
son (1927) and Tixier-Durivault & d’Hondt 
(1974) were based on fragmentary speci- 
mens or were simply listed without descrip- 
tion or comment. In view of the subtle dif- 
ferences among the chrysogorgiid species, 
these records are not accepted until the 
original specimens can be examined. 


VOLUME 114, NUMBER 3 


Fig. 10. Chrysogorgia fewkesii, sclerites from lectotype, MCZ 4850. A, six spindles with flattened tips from 
the body wall region; B, five coenenchymal scales; C, three tentacular rods; D, four pinnular scales. 


VY. 


Material examined.—BL-190, 1, USNM 
49317 (former MCZ 4852); G-190, 1, 
USNM 52789; G-936, 1, USNM 52807; P- 
391, 1, USNM 52845; P-689, 5, USNM 
52848; P-984, 3, USNM 52835; P-1262, 4, 
USNM 52843; specimens reported by 
Deichmann (1936); types of C. fewkesii 
(see below). 

Types.—In his original description, Ver- 
rill (1883: 26) cited several specimens from 
Blake-227, as well as material from “‘sev- 
eral other localities in the same region, and 
off Cuba’’, the Cuban specimen probably 
the one described by Pourtalés (1868) from 
Bibb-22, which Verrill later identified as C. 
fewkesi. However, Deichmann (1936) des- 
ignated the “type” as MCZ 4850, a speci- 
men from BL- 227, which restricts the type 
locality. However, there is more than one 
specimen from the type lot of BL-227; it 
includes 2 colonies, several branches, and a 
slide to which a branch is glued. Thus, one 
of the colonies is chosen as lectotype (MCZ 
4850); the other specimens from this station 
becoming paralectotypes (MCZ 4850a). 
The specimens from Bibb-22 are also con- 
sidered as paralectotypes, but could not be 
found at the MCZ in 2001. No other para- 
lectotypes can be unequivocally determined. 
Type locality as restricted by lectotype: 
13°10'10"N, 61°18'15”"W (off southwestern 
St. Vincent, Lesser Antilles), 1048 m. 

Distribution.—Western Atlantic: Straits 
of Florida, Northwest Providence Channel, 
Jamaica, Lesser Antilles, off Guyana, off 
Colombia; 430—1200 m.? Eastern Atlantic: 
purportedly (see Discussion) from Azores 
and off Morocco; 1022—2165 m (Thomson 
1927, Tixier-Durivault & d’Hondt 1974). 


Chrysogorgia desbonni Duchassaing & 
Michelotti, 1864 
Figs. 11-12 


Chrysogorgia Desbonni Duchassaing & 
Michelotti, 1864: 13 (107), 21 (115), pl. 
1, figs. 7-8, pl. 4, fig. 5.—not Pourtalés, 
1868: 131-132 (=C. fewkesii).—Duchas- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


saing, 1870: 17.—Verrill, 1883: 25-26, 
pl. 2, figs. 6, 6a, 6b. 

Chrysogorgia desbonni: Wright & Studer, 
1889: 24.—Hargitt & Rogers, 1901: 281, 
pl. 1, figs. 1—5.—Versluys, 1902: 85— 
86.—Ktikenthal, 1919: 532; 1924: 
404.—Deichmann, 1936 (in part: not 
specimens from BL-101, BL-297 or BL- 
“XIII” (the latter actually BL-55) and not 
pl. 35, fig. 1, which are C. thyrsiformis), 
228—230, pl. 22, fig. 5, pl. 23, figs. 19— 
27.—Bayer, 1954: 280 (listed); 1958: 389 
(listed). 

Chrysogorgia occidentalis Versluys, 1902: 
56, 86.—Kiikenthal, 1919: 523-524, fig. 
232; 1924: 398. [new synonym] 

Chrysogorgia desbonhi (sic): Grygier, 
1984: 165. 


Description.—Colonies biflabellate, two 
equal-sized, parallel to slightly convex, 
roughly circular flabella originating 15—20 
mm above the base; colonies up to 16 cm 
in height and 12 cm in width. Holdfast en- 
crusting, disc-shaped. Main stem up to 1.6 
mm in diameter. Main stem bronze to dark 
brown; branches paler. Branching sequence 
2/5R, but present only in basal 15—20 mm 
of colony, the two major and several minor 
flabella being uniplanar and dichotomously- 
branched; anastomosis of branches com- 
mon, sometimes producing a fine reticulum. 
Branches extremely closely spaced, only 
about 0.6—1.0 mm apart, producing an or- 
thostiche interval of about 5 mm; only 3 or 
4 orthostiche intervals occur before the dis- 
tal, dichotomously branched flabella origi- 
nate. In large colonies, up to 40 nodes may 
be present in each flabellum, the internodes 
being 3—4 mm in length, resulting in fla- 
bellar branches up to 12—15 cm in length. 
Flabellar branches gradually decrease in di- 
ameter, being quite thick proximally (as 
thick as the main stem) and decreasing to 
about 0.12 mm on terminal twigs, resulting 
in a wiry construction. Dichotomous 
branching usually unequal, one side of each 
dichotomy usually much thicker than the 
other. One polyp, sometimes 2, present on 


VOLUME 114, NUMBER 3 


773 


Fig. 11. 
tall. C, colony from G-391, USNM 52797, height 10 cm, two galatheid crabs attached within flabella. 


each internode, the polyps usually oriented 
such that they project perpendicular to the 
flabella and in an outward direction. Polyps 
small, cylindrical, about 1.5—1.8 mm in 
height. Coenenchymal sclerites elongate, 
warty scales with very irregular margins, up 
to 0.71 mm in length and 0.06 mm in width, 
although most scales are shorter, 1.e., 0.42 
mm in length. Warts of coenenchymal 
scales about 11-17 wm in diameter and 
multiheaded. Body wall sclerites trans- 
versely arranged spindles, often strongly 
curved and somewhat flattened, their distal 
ends being pointed or sometimes flattened 
like a scale. Body wall sclerites up to 0.75 
mm in length and 0.05—0.07 mm in width, 
bearing small, compound warts 7—9 m in 
diameter. Tentacular sclerites straight and 
rod-shaped, up to 0.24 mm in length and 
0.02—0.04 mm in diameter. Pinnular scler- 
ites typical rectangular scales, most 0.11— 
0.14 mm in length and 0.02 mm in width, 
often slightly medially-constricted. 
Discussion.—When a colony is held on 
edge, a space 5-10 mm wide can often be 
seen between the two parallel flabella, or an 
enclosed cavity may be detected in those 
specimens that have convex flabella. In the 
latter case, the flabella resemble the two 


Chrysogorgia desbonni. A—B, neotype, BL-232, MCZ 4839, lateral and edge views, colony 5.5 cm 


halves of a clam, the enclosed space some- 
times providing refuge for galatheid crabs. 
Aplacophoran mollusks are also sometimes 
found attached to the branches of the fla- 
bellum (e.g., O-5419). 

Chrysogorgia desbonni is easily recog- 
nized as the only western Atlantic species 
to have a flabellate colony form (Table 3). 
Comparisons to C. thyrsiformis are made in 
the discussion of that species. 

For reasons not understood, Duchassaing 
& Michelotti (1864) described the genus 
Chrysogorgia and its type species C. des- 
bonni twice independently in the same pub- 
lication, as species 18 and 87. Both speci- 
mens were from Guadeloupe and both de- 
scriptions might even have been based on 
the same specimen; however, species 18 
was reported as 10—13 cm tall, whereas 
species 87 as 8 cm tall. Duchassaing (1870) 
later confirmed that both descriptions per- 
tained to the same species. The relatively 
shallow purported depth at which the spec- 
imen(s) was collected (300—400 m, see Du- 
chassaing & Michelotti 1864: 101) would 
suggest only two possibilities: C. desbonni 
or C. thyrsiformis. The brief French and 
Latin descriptions of the species are not ad- 
equate to distinguish the two, and the types 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 12. Chrysogorgia desbonni. A—D, sclerites from neotype, MCZ 4839. A, five curved spindles from 
body wall; B, five coenenchymal scales; C, two tentacular rods; D, four pinnular scales. E, polyp from neotype, 
width 0.67 mm (from Verrill, unpublished plates). 


VOLUME 114, NUMBER 3 


are lost (Wright & Studer 1889), but the 
figure of the colony of species 87 appears 
to be the biflabellate taxon (1.e., C. desbonni 
as understood in this paper and C. occiden- 
talis sensu Versluys and Kiikenthal), and 
the number of polyps per internode as well 
as the length and number of internodes are 
also consistent with that species. Conse- 
quently a neotype of the biflabellate species 
collected near the type locality has been 
chosen to represent the species (see Types). 

Verrill (1883) also recognized C. desbon- 
ni as a flabellate species, reporting it from 
“numerous” but largely unspecified loca- 
tions in the Caribbean (see Types of C. oc- 
cidentalis, below). Wright & Studer (1889) 
were convinced that Duchassaing & Mich- 
elotti (1864) had described two very differ- 
ent species under the same name, opting to 
recognize the second of the descriptions, 
the flabellate species 87, as the name bearer. 
Hargitt & Rogers (1901) clearly reported 
the biflabellate form as their C. desbonni. 
However, based purely on the literature, 
Versluys (1902) concluded that Verrill’s C. 
desbonni was a different species from the 
one described by Duchassaing & Michel- 
otti, proposing the name C. occidentalis for 
Verrill’s specimens. Versluys did not des- 
ignate type specimens for C. occidentalis 
nor did he actually see any specimens, but 
simply relied on the specimens examined of 
C. desbonni by Verrill, all of which must 
be considered as syntypes of C. occidentalis 
(see ““Types’’). Kiikenthal (1919) followed 
the taxonomy of Versluys and apparently 
examined at least one of Verrill’s specimens 
at the MCZ and illustrated a branch, but he 
still did not have a firm comprehension of 
true C. desbonni. Comparing crude illustra- 
tions and poor descriptions without re- 
course tO examining specimens is ill-ad- 
vised as a basis in taxonomy. But, since the 
types of C. desbonni are lost, the nomen- 
clatural issue may be simply resolved by 
designating a biflabellate specimen from the 
syntype series of C. occidentalis as the lec- 
totype of that species as well as the neotype 
of C. desbonni, making C. occidentalis a 


775 


junior objective synonym of C. desbonni. 
Such a specimen is chosen from BL-232 
(MCZ 4839). 

Because C. desbonni has flattened spin- 
dles in its body wall and tentacles, it be- 
longs to Chrysogorgia (Group “Spiculo- 
sae’’). 

Material examined.—AlIb-2342, 3, 
USNM 44139; Alb-2347, 1, USNM 44140; 
Atl-2980B, 1, MCZ 3866; Azl-2999, 3, 
MCZ 3867; Azl-3326, 9, MCZ 3719-22; 
Atl-3436, 1, MCZ 3705; Atl-3437, 1, MCZ 
3698; Atl-3438, 3, MCZ 3752; Atl-3479, 1, 
MCZ 3756; Caroline-43, 2, USNM 43791; 
Combat-447, 1, USNM 50802; G-232, 1, 
52816; G-235, 1, USNM 52791; G-241, 3, 
USNM 52792; G-242, 1, USNM 52793; G- 
246, 2, USNM 52794; G-261, 1, USNM 
52795; G-387, 1, USNM 52796; G-391, 6, 
USNM 52797; G-678, 2, USNM 52799; G- 
679, 1, USNM 52817; G-692, 2, USNM 
52800; G-798, 1, USNM 52801; G-889, 2, 
USNM 52802; G-897, 1, USNM 52804; G- 
898, 1, USNM 52834 (specimen reported 
by Grygier, 1984); G-899, 1, USNM 52805; 
G-925, 2, USNM 52806; G-927, 1, USNM 
52819; G-936, 2, USNM 52807; G-1125, 
1, USNM 52905; G-1312, 2, USNM 52833; 
G-1314, 1, USNM 52832; Eastward-31281, 
1, USNM 80097; Nutting Iowa Bahama 
Expedition, 1893, 1, off Havana, USNM 
91861; O-4940, 1, USNM 52813; O-10849, 
1, USNM 52837; P-200, 2, USNM 52808; 
P-208, 1, USNM 52824; P-209, 1, USNM 
52809; P-594, 3, USNM 52810; P-610, 5, 
USNM 52811; P-611, 1, USNM 52812; P- 
857, 1, USNM 52826; P-877, 1, USNM 
52827; P-890, 2, USNM 52828; P-931, 1, 
USNM 52830; P-1141, 2, USNM 52836; 
SB-440, 2, USNM 51263; off Sanibel Is- 
land, Florida, depth unknown, 3, USNM 
52814; specimens reported by Verrill 
(1883), Deichmann (1936), and Grygier 
(1984); neotype. 

Types.—The type of C. desbonni is not 
present at the Museo Regionale di Scienze 
Naturali, Torino (Wright & Studer 1889, L. 
Levi, pers. comm., 2001) or the Museum of 
Florence, and thus is presumed to be lost. 


776 


Indeed, none of the specimens reported by 
Duchassaing & Michelotti (1864) have ever 
been found (EK M. Bayer, pers. comm.). To 
resolve the taxonomic confusion about this 
species (see Remarks), a neotype of 5.5 cm 
height is chosen from BL-232 (MCZ 4839). 
Original type locality: Harbor of Moule, 
Guadeloupe, Lesser Antilles (depth 300— 
400 m?). Type locality (as redefined by 
neotype): 13°06’45"N, 61°06'55”W (east of 
St. Vincent, Lesser Antilles), 88 fms (=161 
m). 

The syntypes of C. occidentalis Versluys, 
1902, must be considered to be the material 
reported by Verrill (1883: 26) as C. des- 
bonni, which reads: ““‘numerous localities in 
the Caribbean Sea and among the Antilles, 
in 88 to 163 fathoms, by the Blake, in 
1878-1979, and off Cuba, in 288 fathoms, 
in 1880.” From the station list of the Blake, 
at least three stations correspond to these 
requirements: BL-232 (the shallow end 
range), BL-241 (the deep end range), and 
BL (Bartlett)-V (the Cuban specimen). 
Thirteen additional Blake stations are listed 
by Deichmann (1936), all of which Verrill 
probably examined. Specimens from all of 
these stations are still present at the MCZ, 
except for BL (Bartlett)-V. A lectotype is 
chosen from BL-232 (MCZ 4839), which 
corresponds to one of the extremes of the 
bathymetric range reported by Verrill, and 
certainly one of the specimens he exam- 
ined. Because it is the same specimen cho- 
sen for the neotype of C. desbonni, C. oc- 
cidentalis becomes a junior objective syn- 
onym of C. desbonni. Type Locality: same 
as listed for C. desbonni above. 

Distribution.—Fairly continuous distri- 
bution from off Turneffe, Belize, clockwise 
throughout the Antilles to Grenada, includ- 
ing the Bahamas; 155—595 m. 


Chrysogorgia thyrsiformis Deichmann, 
1936 new rank 
Figs. 13-14 


Chrysogorgia desbonni vat. 
Deichmann, 1936: 230. 


thyrsiformis 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Chrysogorgia desbonni: Deichmann, 1936: 
in part (BL-101, BL-297, BL-S5S5, the latter 
reported as BL-XIID), pl. 35, fig. 1. 

Chrysogorgia elisabethae Bayer, 1951: 
269-272, pl. 9, figs. 56-57; 1952: 189 
[new synonym]; 1954: 280 (listed). 


Description.—Colonies bushy, rarely 
more than 8 cm in height and 8—9 cm in 
width. Holdfast encrusting, disc-shaped. 
Main stem slender (1.1—-1.7 mm in diame- 
ter), straight, and rarely more than 3 cm in 
length; color of main stem golden-brown or 
bronze. Branching sequence 2/5R; branches 
extremely closely spaced, only about 0.5— 
0.7 mm apart, producing an orthostiche in- 
terval of 2.7—3.5 mm on lower part of col- 
ony and about 5 mm near distal portion of 
main stem. First two internodes of each 
branch form a plane that is roughly hori- 
zontal to the substrate G.e., perpendicular to 
the main stem); however, at the second in- 
ternode the orientation of the branch is usu- 
ally twisted 45—90°, the plane of the re- 
maining branch, which usually remains uni- 
planar to the terminal twigs, being vertical 
(i.e., parallel to the main stem). This twist- 
ing of the plane of branching and equal de- 
velopment of branches around the main 
stem produces a bushy colony form. Well- 
developed branches consist of about 10—12 
internodes, each 5—8 mm in length, result- 
ing in branch lengths of up to 75 mm. Di- 
chotomous branching usually results in 
equal-sized sub-branches throughout each 
branch; angle between sub-branches fairly 
small (about 30°) . Branches arise from the 
main stem oriented in an upward direction, 
about 45° from the vertical; branch anasto- 
mosis rare. Branches quite thick near main 
stem but gradually decrease in thickness, 
ending in terminal twigs of 0.12 mm in di- 
ameter; general tensile strength wiry. Two 
to 3 polyps occur on each internode, often 
more on the terminal twigs. Polyps small 
(up to 1.2 mm in height) and often have a 
highly constricted base; they project per- 
pendicular to the branches or sometimes ap- 
pear to angle backward toward the main 


VOLUME 114, NUMBER 3 


V7) 


Fig. 13. 


stem. Sclerites similar to those of C. des- 
bonni. Coenenchymal sclerites elongate, 
warty scales (warts about 8 pm in diameter) 
0.36—0.54 mm in length and 0.05—0.08 mm 
in width, having pointed distal ends and 
highly irregularly-shaped edges. Body wall 
sclerites transversely arranged, curved and/ 
or irregularly-shaped, finely-warty spindles 
up to 0.71 mm in length and 0.06—0.07 mm 
in width. These sclerites are slightly flat- 
tened, particularly at their distal ends, but 
are not interpreted to be scales. Tentacular 
sclerites longitudinally arranged rods simi- 
lar in shape to those of body wall but small- 
er (i.e., 0.21—0.42 mm in length). Pinnular 
sclerites typical rectangular scales 0.18— 
0.21 mm in length. 

Discussion.—Although described as a 
variety of C. desbonni, C. thyrsiformis dif- 
fers from that species in several consistent 
characters, warranting its elevation to the 
species level. C. thyrsiformis differs in hav- 
ing a bushy colony (not flabellate), fewer 
and longer internodes (see Table 3), more 
polyps per internode, slightly shorter and 
flatter body wall spindles, and more nu- 


Chrysogorgia thyrsiformis, stereo view of lectotype, MCZ 4849, height 5.5 cm. 


merous irregularly-shaped spindles in the 
body wall (in addition to the curved, reg- 
ularly-shaped, flattened spindles). As de- 
scribed above, although the branches of C. 
thyrsiformis are basically uniplanar, there is 
often a twisting of the orientation at the sec- 
ond node, and the distalmost 8—12 branches 
that diverge from the main stem produce 
major branches. In contrast, all branches 
that diverge from the main stem of C. des- 
bonni are exclusively uniplanar, only the 
distalmost two forming the two large uni- 
planar flabella having numerous nodes. Be- 
cause C. thyrsiformis has flattened, curved 
spindles in its body wall and tentacles, it 
should, along with C. desbonni, be placed 
in Chrysogorgia (Group Spiculosae). Bayer 
(1951:272) placed C. thyrsiformis (as C. 
elisabethae) in the ““Squamosae aberran- 
tes’’, apparently interpreting the flattened 
tentacular spindles/rods as scales, although 
he clearly noted in his description that both 
body wall and tentacles possessed rods, 
which would place it in Group A. 
Aplacophoran mollusks are sometimes 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 14. Chrysogorgia thyrsiformis. A—D, sclerites of lectotype, MCZ 4849. A, five curved spindles from 
body wall; B, five coenenchymal scales; C, a tentacular rod; D, three pinnular scales. E, polyp of lectotype, 1 
mm in width (from Verrill, unpublished plates). 


VOLUME 114, NUMBER 3 


found attached to branches of this species 
(e.g., G-503). 

Material examined.—Alb-2655, 4, 
USNM 44129; A?/-3320, 1, MCZ; Atl-3480, 
1, MCZ 3670; BL-(Sigsbee), 2 miles east of 
Havana, 1, USNM 49501; Caroline-102, 2, 
USNM_ 100888; R/V Cape_ Florida, 
27°31'N, 79°15'W, 350-400 m, 2, USNM 
73940; CI-37, 2, USNM 54816; G-168, 1, 
USNM 52788; G-233, 10, USNM 52790; 
G-503, 5, USNM 52798; G-705, 1, USNM 
52818; G-893, 1, USNM 52803; G-936, 1, 
USNM_ 100886; Eastward-26549, 7, 
USNM 100889; O-2655, 2, USNM 59807; 
O-5021, 2, USNM 52821; O-5419, 1, 
USNM 53034; P-200, 3, USNM 100887; 
P-739, 6, USNM 52815; P-848, 2, USNM 
5282552-907,.2, USNM 52829; P-991, 1, 
USNM 52831; P-1410, 1, USNM 54815; 
SB-3497, 1, USNM 100890; University of 
Iowa, off Havana, 1, USNM 49489 (re- 
ported by Bayer, 1952); types of C. thyris- 
iformis and C. elisabethae; specimens re- 
ported as C. desbonni by Deichmann 
(1936). 

Types.—Deichmann cited a type of this 
variety from a Hassler station off Barbados 
(MCZ 4849) and another specimen from 
BL-281 (MCZ 4867); however, there are 
two specimens in lot 4849. Thus one spec- 
imen from MCZ 4849 is chosen as the lec- 
totype, the other becoming a paralectotype 
~(MCZ 4849a). The specimen from BL-281 
is also considered to be a paralectotype. 
Type Locality: “‘off Barbados,” 183 m. 

The holotype of C. elisabethae was col- 
lected at Alb-2129, and is deposited at the 
USNM (7552). Type locality: 19°56'04’N, 
75°48'55"W (off Santiago, Cuba), 501 m. 

Distribution.—Fairly continuous distri- 
bution from Grand Bahama Bank through- 
out the Antilles to Isla Tortuga, Venezuela, 
including Yucatan Peninsula (off Cozumel) 
and the Bahamas; 146—526 m. 


Chrysogorgia agassizii (Verrill, 1883) 
Figs. 15-16 


Dasygorgia Agassizii Verrill, 1883: 22-23, 
pl. 2, figs. 4, 4a—b(c—e); 1884: 220; 1885: 


TI) 


511-512, pl. 9, fig. 199.—Agassiz, 1888: 
143-144, fig. 455.—?Roule, 1896: 304— 
305. 

Chrysogorgia agassizii: Versluys, 1902: 
60.—not Nutting, 1912: 55—56.—Ktiken- 
thal, 1919: 530, fig. 233; 1924: 403.— 
Deichmann, 1936: 222, 233-234, pl. 23, 
figs. 34—40, but not pl. 34, figs. 1-2 (=C. 
elegans).—Madsen, 1944: 49-54, figs. 
42—47.—?Tyler & Zibrowius, 1992: 217. 

Chrysogorgia agassizi: Kinoshita, 1913: 3 
(listed).—?Tixier-Durivault & d’Hondt, 
1974: 1409.—?Grasshoff, 1982a: 748 
(map 4); 1982b: 949, figs. 22—24.—Vin- 
ogradov, 2000: 101-103, figs. 1-2. 

?Chrysogorgia pentasticha: Thomson, 
NODS QD, 

?Chryzogorgia (sic) agassizi: Pasternak, 
kes SY, 


Description.—Colony bottlebrush- 
shaped, up to 41 cm in height, and attached 
by an encrusting or rhizoidal holdfast. Main 
stem up to 2.5 mm in diameter; golden- 
brown in luster. Branching sequence con- 
sistently 2/5R: branches closely spaced, one 
every |.5—2.0 mm; orthostiche interval 10— 
12 mm; branches diverge from main stem 
at about 70° from the vertical. Terminal 
twig branch diameter quite small (0.1 mm), 
producing a flexible, limp colony. Number 
of nodes per branch ranges from 2—5, but 
are usually 3, the first 2 internodes being 
short (2—4 mm), the third 4—6 mm, and the 
terminal twigs being up to 50 mm in length, 
combining to produce a total branch length 
of 60—65 mm. Branching dichotomy not al- 
ways symmetrical, the internodes and ter- 
minal twigs of a branch usually arranged in 
a horizontal plane. Polyps usually absent 
from first 2 internodes, one occurring near 
the third internode, and up to 11 polyps on 
the terminal twigs. Polyps cylindrical, up to 
2.5 mm in height and about 0.75 mm in 
diameter, obliquely arranged on the branch- 
es. Coenenchymal sclerites smooth, elon- 
gate scales with rounded distal ends and 
finely serrate edges, up to 0.36 mm in 
length and 0.04 mm in width. Body wall 


780 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 15. 
MCZ 5748, height 10.5 cm. 


sclerites similar to coenenchymal sclerites 
but slightly larger: 0.42—0.48 mm in length 
and 0.06—0.07 mm in width. Body wall 
scales longitudinally arranged, with smooth 
to finely granular (but not warted) faces and 
finely serrate edges. Bluntly-tipped, finely- 
granular, straight rods 0.42—0.60 mm in 
length and 0.08—0.13 mm in width occur at 
the base of the tentacles, but are quite rare, 
perhaps occurring with a frequency of one 
per tentacle. Tentacular rods bear small 
compound warts, each about 8 wm in di- 
ameter, which are composed of smaller el- 
ements. Pinnular sclerites smooth elongate 
scales up to 0.21 mm in length and 0.04 
mm in width, having finely-serrate edges. 
Discussion.—This species is exhaustive- 
ly described, figured, and discussed by 
Madsen (1944), based on several specimens 
collected south of Iceland; however, the 
eastern Atlantic records of C. agassizii (see 
*“‘Distribution’’) have been little more than 
unfigured and undescribed listings of the 


Chrysogorgia agassizii. A, lectotype, BL-308, MCZ 4870, height 15 cm; B, paralectotype, BL-308, 


species. Since I have not examined them, 
their synonymy records are queried. 

It is of interest to note that the specimens 
from the four Albatross stations reported 
herein were originally identified but not re- 
ported by A. E. Verrill. 

Based on repeated photographs of a spec- 
imen attached to the bow rail of the R. M. 
S. Titanic, Vinogradov (2000) calculated 
the growth rate of this species to be about 
1 cm per year. 

Material examined.—Alb-2034, 1, 
USNM 9150; Alb-2035, 4, USNM 17253; 
Alb-2220, 1, USNM 7920; Alb-2573, 2, 
USNM 11854; Knorr 58-I-827, 1, USNM 
59808; Gyre MA2:04, 1, USNM 88109; 
Nutting’s (1912) misidentification, Alb- 
5080, USNM 30170; types of D. agassizii 
(see below). 

Types.—The larger of the two syntypes is 
deposited at the MCZ (4870) and a smaller 
one is at the USNM (5748). Deichmann 
(1936), in designating MCZ specimen 4870 


VOLUME 114, NUMBER 3 


* 


ta. 


Rrra ens 


Chrysogorgia agassizii, sclerites from paralectotype, BL-308, MCZ 5748. A, five scales from body 


Fig. 16. 
wall; B, five coenenchymal scales; C, enlargement of tip of tentacular rod; D, four tentacular rods; E, four 


pinnular scales. 


Fig. 17. Chrysogorgia squamata, \ectotype, BL- 
283, MCZ 4862, height of colony 10.5 cm. 


as the type, effectively makes it the chosen 
lectotype for the species, the USNM spec- 
imen thus becoming a paralectotype. Type 
Locality: Blake-308: 41°04'45"N, 65°35’ 
30”"W (off Georges Bank), 2271 m. 

Distribution.—Western Atlantic: off con- 
tinental slope of northeastern United States 
and Newfoundland from 39—42° N latitude, 
1928-3700 m. North and eastern Atlantic: 
off Iceland (Madsen 1944);? west of Ireland 
(Tyler & Zibrowius 1992);? Bay of Biscay 
(Roule1896, Grasshoff 1982a);? Azores 
(Thomson 1927);? Atlantis Seamount (Pas- 
ternak 1985); 1425-2860 m. 


Chrysogorgia squamata (Verrill, 1883) 
Figs. 17-18 


Dasygorgia squamata Verrill, 1883: 24. 

Chrysogorgia squamata: Versluys, 1902: 
85.—Ktkenthal, 1919: 538; 1924: 
408.—Deichmann, 1936: 232-233, pl. 
22, fig. 8, pl. 23, figs. 28-33. 


Description.—Colonies bottlebrush- 
shaped, up to 20 cm in height and 6 cm in 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


width, attached by an encrusting holdfast. 
Main stem up to 1.6 mm in diameter, yel- 
lowish-amber in luster, and, along with the 
internodes, covered with numerous, slender 
cnidal papillae up to 0.2 mm in height. 
Branching sequence predominantly 1/5R, 
although the occasionally the sequence 
shifts to 1/6 or even 1/7R. Branches occur 
every 2.1—3.0 mm, resulting in an ortho- 
stiche interval for the 1/5 sequence of 8.5— 
12.0 mm. Branches diverge from main stem 
at 70—90° from the vertical. Distal branch 
diameter 0.25 mm; branches and colony 
wiry in construction. Number of nodes per 
branch 2 or 3. First internode 4—8 mm in 
length; second, 4—5 mm; third, 4—5 mm, 
and terminal twigs up to 10 mm, resulting 
in a total branch length up to 28 mm. Pol- 
yps sparse, usually absent from the first 2 
internodes, only 1 on the third internode, 
and | or 2 on the terminal twigs. Polyps 
cylindrical, up to 1.8 mm in height. Coe- 
nenchymal sclerites elongate (up to 0.45 
mm in length and about 0.07 mm in width), 
smooth scales, rounded on their distal ends, 
with finely serrate edges, and often slightly 
constricted medially. Body wall sclerites 
transversely arranged, elongate (up to 0.54 
mm in length and 0.08—0.09 mm in width), 
smooth scales, also with rounded distal 
ends and finely serrate edges, the apices of 
the serrations of uniform height (1—2 pm) 
and quite regularly spaced (every 3—4 wm). 
Backs of tentacles, adjacent to the naked 
space at the base of each tentacle, filled 
with longitudinally-placed, warty rods, up 
to 0.42 mm in length and 0.07 mm in di- 
ameter. Smaller rods (0.20—0.25 X 0.06 
mm) occur in the distal tentacles. Pinnular 
scales quite small, somewhat rectangular 
and usually slightly medially constricted. 
Pinnular scales up to 0.17 mm in length and 
0.09 mm in width, but may be much small- 
er, e.g., 0.08 XK 0.02 mm. 
Discussion.—Deichmann’s (1936) rede- 
scription of this species, which must have 
been based on the type material, was some- 
what misleading, in that she indicated a 
branching sequence of 1/7, branches with 


VOLUME 114, NUMBER 3 


yy 
ty: 


4) pe 
ba St) Sd 


Fig. 18. Chrysogorgia squamata, sclerites from lectotype, MCZ 4862. A, six scales from body wall; B, four 
coenenchymal scales; C, three tentacular rods; D, four pinnular scales. 


784 


5—6 nodes, and longitudinally arranged 
body wall sclerites. As reported in the de- 
scription, the branching sequence is pre- 
dominantly 1/5, branches usually have 2—3 
nodes, (never as many as 6), and body wall 
sclerites are transversely arranged. Versluys 
(1902) tentatively placed C. squamata in 
the group ““Squamosae typicae”, but ac- 
knowledged that he did not have knowledge 
of the tentacular sclerites. Since the tenta- 
cles of this species contain well developed 
rods, it thus belongs in the group ““Squa- 
mosae aberrantes”’. 

Chrysogorgia squamata is a very rarely 
collected species, the specimens listed be- 
low being the first new records since its 
original description. It is easily distin- 
guished from other western Atlantic species 
by having a 1/5R branching sequence and 
cnidal papillae. Indeed, no other species in 
the ““Squamosae aberrantes”’ group has a 1/ 
5R branching sequence (Table 1). 

Material examined.—Atl-3326, 1, MCZ 
(unnumbered); P-607, 1, USNM 52840; 
types of D. squamata from BL-283 (see be- 
low). 

Types.—Verrill reported three specimens 
from two Blake stations, which must be 
considered as syntypes: one from BL-227 
and two from BL-283 (MCZ 4862). Deich- 
mann (1936) restricted the type locality to 
BL-283, but did not designate a lectotype. 
Thus, one of the three (not two) specimens 
from BL-283 (MCZ 4682) is herein desig- 
nated as the lectotype: not the largest col- 
ony of 20 cm height, but the smaller of 12 
cm height that is attached to the substrate. 
The remaining two specimens from BL-283 
(MCZ 4682a) and that from BL-227 are 
thus paralectotypes; however, the latter 
specimen could not be found at the MCZ 
nt ZOOM iyvipe localiinvaaels.0sd05mNr 
59°40'50"W (off southwestern Barbados, 
Lesser Antilles), 431 m. 

Distribution.—off Banco Chinchorro, 
Yucatan Peninsula; southwestern Cuba; 
Lesser Antilles (Barbados and St. Vincent); 
431-1046 m. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Key to the western Atlantic species of 
Chrysogorgia 


The two previous keys (Deichmann 
1936; Bayer 1951) to the western Atlantic 
species of Chrysogorgia relied on the po- 
sition and/or kind of sclerites in the body 
wall as the first couplet. The following key 
emphasizes the more gross and easily rec- 
ognized characters such as colony shape 
and branching sequence, reserving charac- 
teristics of sclerite position, shape and size 
for the terminal couplets. 

1. Colones bottlebrush-shaped; distance 
between branches over 1 mm ........ 2 
1’. Colonies bushy or flabellate; distance 


between branches less than 1 mm ..... 8 

2. Sclerites of body wall longitudinally ar- 
ranged .2 26 aise | ee ee ee 3 

2'. Sclerites of body wall transversely ar- 
ranged. 2436. Sosa eee 6 
. Branching sequence 2/5R ............ 4 
3’. Branching sequence 3/8R _—_C. herdendorfi, 

n. sp. 

4. Body wall sclerites exclusively scales 
Sond den & bic See ee to C. agassizii 

4’. Body wall sclerites includes spindles 
and/or rods ....... 4. - 32 ) 


5. Number of nodes per branch O—2, usu- 

ally 1; distance between branches 2.5— 

3.0 mm C. herdendorfi, n. sp. 
5’. Number of nodes per branch 1—3, usu- 

ally 2; distance between branches 1.0— 

1.5 mm. C. elegans 
5”. Number of nodes per branch 3—6, usu- 

ally 4; distance between branches 3.5— 

ASS WN si. a wae eee C. spiculosa 
6. Branching sequence 2/5R ............ oh 
6’. Branching sequence 1/5R ... C. squamata 
7. Body wall sclerites cylindrical spindles 

up to 0.06 mm in width ...... C. fewkesii 
7’. Body wall sclerites flattened spindles up 

to 0.13 mm in width C. multiflora 
8. Colony bushy; up to 12 internodes per 

branchi.. {5 3k: Sas See C. thyrsiformis 
8’. Colony flabellate (biflabellate); up to 40 

internodes per branch ....... C. desbonni 


Acknowledgments 


I am indebted to Frederick M. Bayer, 
who suggested doing this project, and who 


VOLUME 114, NUMBER 3 


provided help and encouragement through- 
out. I thank Charles (“‘Eddie”’) Herdendorf, 
who collected the type material of the new- 
ly described species and made the loan of 
those specimens possible. I also thank Ardis 
Johnson (MCZ) and Sheila Halsey (BM), 
who generously extended to me the use of 
their collections and facilities or loaned me 
specimens used in this study. Finally, I 
thank Rosemarie Baron-Szabo for her help- 
ful German translations, and Linda Cole, 
for invaluable technical support. 


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Expedition 5(13):65 pp., 1 pl., 55 figs. 

Nutting, C. C. 1908. Descriptions of the Alcyonaria 
collected by the U. S. Bureau of Fisheries 
steamer Albatross in the vicinity of the Hawai- 
ian Islands in 1902.—Proceedings of the United 
States National Museum 34:543—601, pls. 41— 
Slt. 

. 1912. Descriptions of the Alcyonaria collect- 

ed by the U. S. Fisheries steamer “Albatross,” 

mainly in Japanese waters, during 1906.—Pro- 
ceedings of the United States National Museum 

43:104 pp., 21 pls. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Pasternak, EF A. 1985. Specific composition and the 
ways of forming of the bottom fauna isolated 
underwater rises. Gorgonarians and antipatari- 
ans (sic) of the seamounts Rockeway (sic), At- 
lantis, Plato, Great-Meteor and Josephin (sic) 
(Atlantic Ocean).—Trudy Institute of Oceanol- 
ogy 120:21—38, 4 figs. [in Russian]. 

Pourtalés, L. EK 1868. Contributions to the fauna of the 
Gulf Stream at great depths (2nd series).—Bul- 
letin of the Museum of Comparative Zoology 
1(7):121-142. 

Roule, L. 1896. Résultats scientifiques de la campagne 
du “Caudan” dans le Golfe de Gascogne -Aout- 
Septembre 1895. Coelentérés.—Annales de 
l’Université de Lyon 26:299—323. 

Studer, T. 1887. Vesuch eines Systemes der Alcyon- 
aria.—Archiv ftir Naturgeschichte 53(1):74 pp., 
1 pl. 

Thomson, J. A. 1927. Alcyonaires provenant des cam- 
pagnes scientifiques du prince Albert Ier de 
Monaco.—Résultats des Campagnes Scienti- 
fiques accomplies sur son yacht par Albert ler, 
Monaco 73:77 pp., 6 pls. 

, & W. D. Henderson. 1906. An account of the 
Alcyonarians collected by the Royal Indian Sur- 
vey Ship Investigator in the Indian Ocean. Part 
1. The Alcyonarians of the Deep Sea. xvi + 132 
pp-, 10 pls. Indian Museum, Calcutta. 

Tixier-Durivault, A., & M. J. d Hondt. 1974. Les Oc- 
tocoralliaires de la campagne Biagores.—Bul- 
letin du Muséum National d’ Histoire Naturelle, 
Zoologie (3)174(252):1361—-1433, 32 figs. 

Tyler, P. A., & H. Zibrowius. 1992. Submersible ob- 
servations of the invertebrate fauna on the con- 
tinental slope southwest of Ireland (NE Atlantic 
Ocean).—Oceanologica Acta 15(2):211—226, 6 
figs. 

Verrill, A. E. 1883. Report on the Anthozoa, and on 
some additional species dredged by the 
“Blake” in 1877-1879, and by the U. S. Fish 
Commission Steamer “Fish Hawk” in 1880— 
82.—Bulletin of the Museum of Comparative 
Zoology Harvard 11:72 pp., 8 pls. 

. 1884. Notice of the remarkable marine fauna 

occupying the outer banks off the southern 

coast of New England, No. 9. Brief contribu- 
tions to zoology from the Museum of Yale Col- 
lege. No. 55.—American Journal of Science 

(3)28:213-220. 

. 1885. Results from the explorations made by 

the steamer Albatross off the northern coast of 

the United States in 1883.—Annual Report of 
the Commissioner of Fish and Fisheries for 

1883, 503-699, 44 pls. 

. unpublished. The Alcyonaria of the “Blake” 
Expeditions. A. E. Verrill’s unpublished 
plates —140 unpublished plates. 

Versluys, J. 1902. Die Gorgoniden der Siboga-Expe- 


VOLUME 114, NUMBER 3 


dition. I. Die Chrysogorgiiden.—Siboga-Expe- 
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Wright, E. P, & T: Studer. 1889. Report on the Alcyonaria 
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):788—793. 2001. 


Reevaluation of Tropidopathes saliciformis Silberfeld: A hydroid 
originally identified as an antipatharian coral 


Dennis M. Opresko and Rosemarie C. Baron-Szabo 


(DMO) Oak Ridge National Laboratory, Oak Ridge, Tennessee, 37830, U.S.A.; 
(RCB-S) Institute of Paleontology, University of Erlangen, Erlangen, Germany 


Abstract.—Tropidopathes saliciformis Silberfeld (1909) was originally iden- 
tified as an antipatharian coral growing over a hydroid colony. A re-exami- 
nation of the type specimen revealed no evidence of any antipatharian skeletal 
material on the hydroid, and it is assumed that the lower parts of the hydrocauli 
of the hydroid, which lacked hydrocladia, had been mistaken for antipatharian 
sclerenchyme. The hydroid was subsequently identified by Stechow (1913) as 
Halicornaria ishikawai Stechow, 1907 (now known as Gymnangium ishika- 
wai). Therefore, the genus Tropidopathes is herein removed from the Antipa- 
tharia, and for nomenclatural purposes, Tropidopathes saliciformis Silberfeld 
is considered a junior subjective synonym of Gymnangium ishikawai (Ste- 


chow). 


In 1909, Silberfeld described a new ge- 
nus and new species of antipatharian coral, 
Tropidopathes saliciformis. She reported 
that it was growing over a hydroid colony, 
and because of the epizotic habit, she re- 
ferred the genus to the subtribe Crustosae 
which had been established by Schultze 
(1896) for Savagliopsis pedata (Gray). 
Brook (1889) had reported that the lower 
parts of the corallum of the type specimen 
of Antipathes pedata Gray appeared to be 
growing Over a gorgonian axis, and Schul- 
tze (1896) considered this sufficient reason 
for establishing a new genus for the species. 
(NOTE: recent re-examination of the type 
specimen of A. pedata deposited in the Nat- 
ural History Museum, London, revealed no 
evidence of an epizotic habit). In Schultze’s 
revised classification of the Antipatharia, 
the subtribe Crustosae was placed in an un- 
named tribe characterized by the lack of 
peristomal folds; the tribe was placed in the 
subfamily Dekamerota (characterized by 
polyps with ten septa); and the subfamily 
was placed in the family Antipathidae. Van 
Pesch (1914) applied the name Apuchae- 
phora to Schultze’s un-named tribe contain- 


ing the Crustosae. Pax (1918, 1987) did not 
recognize Schultze’s subdivisions of the 
family and, although he retained Tropido- 
pathes as one of seven genera in the Anti- 
pathidae, he noted that the systematic po- 
sition of the genus was unclear. There is no 
evidence that any later antipatharian worker 
had re-examined the type specimen of Tro- 
pidopathes saliciformis after Silberfeld’s 
original description appeared in 1909. 

In Silberfeld’s description of Tropidopa- 
thes saliciformis, she mentions that the 
branches of the antipatharian arise singly 
from a reticulum (““‘Geflecht’’) which is also 
overgrown by bryozoans, sponges, and bal- 
anids. She reported that the branches were 
light brown in color, reached a maximum 
length of 14 cm, and that each had two 
rows of very thick spines. The spines were 
described as being 285 jm wide at the base, 
up to 357 wm in height, rounded at the 
apex, and up to 535 wm apart in the same 
row. She further reported that on the side 
opposite the spines, there was a long crest- 
like ridge, 178 wm in thickness. The illus- 
trations given by Silberfeld (1909, figs. 3— 
4) showed protuberances on either side of 


VOLUME 114, NUMBER 3 


the stem which were arranged bilaterally 
and alternately. These protuberances dif- 
fered from typical antipatharian spines in 
that they appeared slightly concave on one 
side. 

Re-examination of the type specimen.— 
Figure 1 shows the complete holotype of 
Tropidopathes saliciformis Silberfeld in the 
collections of the Zoologische Staatssamm- 
lung Muenchen. Numerous unbranched 
““stems’’ arise from a tangled hydrorhizal 
mass. At the top of some of these “‘stems”’ 
there are small pinnately branched struc- 
tures that Silberfeld had identified as a hy- 
droid (Fig. 2A). A close examination of one 
stem (Fig. 2B) reveals that Silberfeld had 
mistakenly identified the apophyses of the 
hydrocauli of the hydroid as antipatharian 
spines. Associated with each apophysis are 
three openings on the hydrocaulus corre- 
sponding to cauline nematothecae. The hy- 
droid was subsequently identified by Ste- 
chow (1913) as Halicornaria ishikawai Ste- 
chow, 1907. A complete redescription of 
the hydroid is given below. 


Systematic Treatment 


Plumularioidea McCrady, 1859 
Aglaopheniidae Marktanner-Turneretscher, 
1890 
Gymnangiinae Calder, 1997 
Gymnangium ishikawai (Stechow, 1907) 
Figs. 1-3 


2?Aglaophenia balei Marktanner-Turner- 
etscher, 1890:272, pl. 7, figs. 19—20 (sen- 
su Ritchie, 1910:22—23, pl. 4, fig. 12 (see 
Stechow, 1913). 

?Halicornaria flava Nutting, 1905:955, pl. 
13, figs. 11-12 (see Stechow, 1913). 

Halicornaria ishikawai Stechow, 1907: 
198; 1909:100—101; 1913:95. 

Tropidopathes saliciformis Silberfeld, 
1909:19—20, figs 3—4. 

Gymnangium ishikawai.-Stechow, 1923:19; 
Yamada, 1959:84; Hirohito, 1995:290. 


Material examined.—Zoologische 
Staatssammlung Muenchen, No. 12la of 


789 


the collection of Doflein, Japan, Sagami 
Bay, near Jogashima Island, 150 m, 31 Oct 
1904; holotype of Tropidopathes salicifor- 
mis Silberfeld (1909) (schizoholotype, 
USNM 100479). 
Description.—Unbranched monosiphon- 
ic hydrocauli arising from complex root- 
like hydrorhiza (Fig. 1). Hydrocauli up to 
14 cm long and up to 520 pm in diameter; 
separated into internodes about 900 wm in 
length by faint diagonal constrictions. Each 
internode with two frontal lateral hydrocla- 
dial apophyses (Fig. 2). Apophyses distally 
inclined, alternately arranged; surrounded at 
base by three cauline nematothecae, one 
median inferior and two axillary (Fig. 2B). 
Frontal axillary nematotheca very wide, 
leaf-like. Longitudinal crest extending 
down hydrocauli on side opposite apophy- 
ses. Hydrocladia present at top of some, but 
not all hydrocauli. Hydrocladia simple, in 
two rows, bilaterally arranged. Hydrothecae 
in single series on inner side of hydrocladia 
(Fig. 3A). Upper part of hydrotheca tube- 
shaped in lateral view (Fig. 3B); margins 
flared outward, usually forming a single 
weak cusp on each lateral side; adcauline 
margin straight or slightly concave: abcau- 
line margin convex. Hydrothecal aperture 
approximately 160 1m in diameter from ad- 
cauline to abcauline wall. Each hydrotheca 
with three nematothecae; one median infe- 
rior, and two lateral (Fig. 3A, B). Median 
inferior nematotheca tube-like in shape, 
narrowing at apex; lateral nematothecae 
bowl-shaped and relatively wide at apex. 
Terminal aperture of median inferior ne- 
matotheca facing distally; apertures of lat- 
eral nematothecae facing distally and some- 
what medially (i.e., towards each other). 
Distal side of median inferior nematotheca 
mostly free, not adnate to the abcauline 
wall of hydrotheca, and not extending be- 
yond abcauline margin of hydrotheca. Ab- 
cauline intrathecal septum extending about 
a third of the way across interior cavity of 
hydrotheca. Distinct crest-like ridge extend- 
ing down the side of the hydrocladia di- 


790 


o 
4 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 
proximately 15 cm tall. 


rectly opposite side on which hydrothecae 
occur. Gonosomes not present. 
Remarks.—The length of the hydrocauli 
(maximum 14 cm) and the size, shape and 
spacing of the apophyses match very close- 
ly the description given by Silberfeld of the 
‘“‘antipatharian’’ Tropidopathes salicifor- 
mis, and there is little doubt that Silberfeld 
was describing the lower parts of the hy- 
droid colony as an antipatharian. In 1913 
Stechow identified the hydroid as Halicor- 


Gymnangium ishikawai (Stechow). Holotype of Tropidopathes saliciformis Silberfeld (1909); ap- 


naria ishikawai Stechow (1907), and listed 
as the ““‘Untergrund”’ or substrate, the anti- 
patharian Tropidopathes saliciformis, in 
contradiction to Silberfeld’s description of 
the hydroid being the substrate for the an- 
tipatharian. Stechow apparently did not re- 
view Silberfeld’s original description. 
Comparisons.—When Stechow (1913) 
identified Silberfeld’s specimen as Halicor- 
naria ishikawai (Stechow), he noted that 
the median inferior nematothecae were sig- 


VOLUME 114, NUMBER 3 


791 


Fig. 2. 


Gymnangium ishikawai (Stechow); schizoholotype of Tropidopathes saliciformis Silberfeld (1909), 


USNM 100479. A, Upper part of hydrocaulus showing basal part of one hydrocladia. B, Enlarged view of 


hydrocaulus. 


nificantly shorter than those in the type 
specimen of A. ishikawai that he had illus- 
trated in 1909. Because of this, Stechow 
(1913) suspected that the specimen might 
be referrable to Aglaophenia balei Marktan- 
ner-Turneretscher, 1890, as described by 
Ritchie (1910), or to Halicornaria flava 
Nutting, 1905. Stechow (1913), however, 
also mentioned that the margin of the hy- 
drotheca in H. ishikawai was less serrated 
than in either of these two species; in par- 
ticular, he noted that there was no denticle 
on the front or back central areas as de- 
scribed by Nutting. 

Gymnangium balei (Marktanner-Turner- 
etscher) was synonomized with G. hians 
(Busk) by Vervoort & Vasseur (1977). In 
describing a specimen of G. hians, Rees & 
Vervoort (1987) note that the length of the 
medial nematotheca varied considerably de- 


pending on its location on the hydrocladium 
(usually longer on the part of the hydro- 
cladium closest to the hydrocaulus). In the 
specimen of G. hians described by Vervoort 
& Vasseur (1977), the hydrothecal margin 
was reported to have only a single cusp on 
either side, similar to the situation in G. ish- 
ikawai; however, in the specimen of G. 
hians described by Rees & Vervoort 
(1987), the hydrothecal margin was report- 
ed to have three cusps on either side. As 
noted by Calder (1997), not only the length 
of the median inferior nematotheca, but also 
the dentition of the hydrothecal margin are 
quite variable in species of Gymnangium. 
Hirohito (1995) reported and illustrated ma- 
terial of G. ishikawai and G. hians from 
Sagami Bay, Japan. Both species were de- 
scribed as having median inferior nemato- 
thecae extending beyond the hydrothecal 


Fig. 3. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


pa 
on] 


a 


‘wit 


Gymnangium ishikawai (Stechow); upper part of the schizoholotype of Tropidopathes saliciformis 


Silberfeld (1909), USNM 100479. A, Lower portion of hydrocladium, frontal view with two hydrotheca. B, 
Lateral view of hydrocladium showing a single hydrotheca and associated nematotheca. 


margin; differentiated mainly on the basis 
of the serration of the hydrothecal margin 
(inconspicuous in G. ishikawai, and very 
conspicuous in G. hians). According to Cal- 
der (personal communication), G. ishikawai 
is distinct from G. hians Busk. In the type 
material of Tropidopathes saliciformis the 
dentition along the hydrothecal margin is 
relatively weak, suggesting that Silberfeld’s 
specimen should be referred to G. ishika- 
wal, however, the median inferior nemato- 
theca is relatively short and its adcauline 
side is not adnate to the abcauline hydro- 
thecal body wall, suggesting that Silbereld’s 
specimen may be distinct from both G. ish- 
ikawai and G. hians. 


Acknowledgments 


The authors wish to thank B. Ruthenstein 
of the Zoologische Staatssammlung 
Muenchen for his kind hospitality, and for 
making the holotype of 7ropidopathes sal- 
iciformis Silberfeld available for study. We 
also wish to thank S. Braden of the Smith- 
sonian Institution for preparing the speci- 


mens for scanning electron micrography. 
Special thanks to S. Cairns of the Smith- 
sonian Institution and to D. Calder of the 
Royal Ontario Museum for their careful re- 
views of the manuscript, and to S. Gardiner 
of Bryn Mawr College for his editorial re- 
view. This work was supported in part by 
the Smithsonian Institution, Washington, 
D.C., and in part, by Oak Ridge National 
Laboratory, Oak Ridge TN. 


Literature Cited 


Brook, G. 1889. Report on the Antipatharia—Reports 
of the Scientific Results of the Voyage of the 
H.M.S. Challenger, Zoology 32:5—222. 

Busk, G. 1852. An account of the Sertularia of the 
Rattlesnake. Macgillivray’s Narrative of the 
Voyage of the Rattlesnake. 1, Appendix 4, pp. 
385—402. 

Calder, D. R., 1997. Shallow-water hydroids of Ber- 
muda: Superfamily Plumularioidea.—Royal 
Ontario Museum, Life Sciences Contributions, 
No. 161, pp. 1-82. 

Marktanner-Turneretscher, G. 1890. Die Hydroiden des 
Naturhisorischen Hofmuseums.—Annalen des 
Hofmuseums, Wien. 5, pp. S-195—286. (Not 
seen) 


VOLUME 114, NUMBER 3 


McCrady, J. 1859. Gymnopthalmata of Charleston 
Harbor.—Proceedings of the Elliott Society of 
Natural History 1:103—221. 

Nutting, C. 1905. Hydroids of the Hawaiian Islands 
collected by the Steamer Albatross in 1902.— 
Bulletin of the U.S. Fish Commission for 1903, 
pp. 931-959. 

Pax, E 1918. Die Antipatharien.—Zoologische Jahr- 
bticher 41:419—478. 

. 1987. Ordre des Antipathaires (revised by M. 
Van-Praét and D. Doumenc). Pp. 189-210 in 
Traité de Zoologie: Anatomie, Systématique, 
Biologie, ed., P-P. Grassé, Tome 3, Cnidaires 
Anthozoaires, Fasc. 3. D. Doumenc, ed., Ma- 
son, Paris. 

Rees, W. J., & W. Vervoort. 1987. Hydroids from the 
John Murray Expedition to the Indian Ocean, 
with revisionary notes on Hydrodendron, Abie- 
tinella, Cryptolaria and Zygophylax (Cnidaria: 
Hydrozoa).—Zoologische Verhandelingen 237: 
1—209. 

Ritchie, J. 1910. Hydroids of the Indian Museum.— 
Records of the Indian Museum 5(1):1—30. (Not 
seen) 

Schultze, L. S. 1896. Beitrag zur Systematik der An- 
tipatharien.—Abhandlungen der Senckenber- 
gischen naturforschenden Gesellschaft 23:1—40. 

Silberfeld, E. 1909. Japanische Antipatharien.—Ab- 


193 


handlungen der mathematisch-physikalischen 
Klasse der Ko6niglichen bayerischen Akademie 
der Wissenschaften Supplement 1, vol. 7, pp. 
1-30. 

Stechow, E. 1907. Neue japanische Athecata and Plu- 
mularidae aus der Sammlung Dr. Doflein.— 
Zoologischer Anzeiger 32:192—200. 

. 1909. Hydroidpolypen der Japanische Ostkus- 

te. I. Teil, Athecata und Plumularidae.—Abhan- 

dlungen der mathematisch-physikalischen Klas- 
se der K6niglichen bayerischen Akademie der 

Wissenschaften. Supplement Band 1, Abhan- 

dlung 2, pp. 1-162. 

. 1913. Hydroidpolypen der Japanischen Ost- 

kuste. If. Teil, Athecata und Plumularidae.— 

Abhandlungen der mathematisch-physikalisch- 

en Klasse der K6niglichen bayerischen Akade- 

mie der Wissenschaften Supplement Band 3, 

Abhandlung 6, pp. 1-111. 

. 1923. Die Hydroidfauna der Japanische Re- 
gion.—Journal of the College Science, Tokyo 
Imperial University, 44(8):1—23. 

van Pesch, A. J. 1914. The Antipatharia of the Siboga 
Expedition.—Siboga-Expeditie Monographe 
17:1—258. 

Veevoort W., & P. Vasseur. 1977. Hydroids from 
French Polynesia with notes on distribution and 
ecology.—Zoologische Verhandelingen 159:1— 
98. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):794—821. 2001. 


East Pacific Mexican Tethya (Porifera: Demospongiae) with 
descriptions of five new species 


Michele Sara, Patricia GOmez, and Antonio Sara 


(MS, AS) Dipartimento per lo Studio del Territorio e delle sue Risorse, Universita di Genova, 
Corso Europa 26, 16132 Genova, Italy 
(PG) Instituto de Ciencias del Mar y Limnologia, U. N. A. M. Circuito Exterior S/N, 
Ciudad Universitaria, C.P. 04510, México, D.E 


Abstract.—Seven species of Tethyidae (Porifera: Demospongiae: Hadrom- 
erida) inhabiting the Pacific waters of Mexico, including the Gulf of California, 
were studied. Five of them, Tethya ensis, T. mexicana, T. ovum, T. paroxeata, 
and 7. socius are new species. The remaining two, T. taboga De Laubenfels 
and 7. californiana De Laubenfels, are new records for the area. 


This is the first published report on spe- 
cies of the genus Tethya (Porifera: Demos- 
pongiae) coming from the Pacific coast of 
Mexico. A total of 242 collected Tethya 
specimens yielded five new species and two 
other species recorded for the first time. 
Tethya taboga (De Laubenfels) was known 
only for the Pacific coast of Panama, and 
T. californiana De Laubenfels, 1932, rede- 
scribed by Sara & Corriero (1993), is re- 
corded here for a more southern site in 
Mexico. Only four species of Tethya were 
previously known for the whole Pacific 
coast of America: T. californiana De Lau- 
benfels from California, 7. taboga (De Lau- 
benfels) from Panama, 7. papillosa Thiele 
from Calbuco (Chile) and 7. sarai Des- 
queyroux-Fatnndez & van Soest, 1997 from 
the Galapagos Islands. This small number 
of American Pacific species, now amount- 
ing to nine, is probably due both to a real 
scarcity of species and to the poverty of re- 
cords from this area. Donatia multifida 
Carter, 1882 from Acapulco (Mexico), ex- 
amined by Sara (1994) on spicular slides of 
the British Museum of Natural History 
(BMNH) is currently regarded as incertae 
sedis, given the scarcity of the material. 
Moreover, it is impossible to know if it be- 
longs to a species of the Tethya seychellen- 


sis group or if it should be considered as 
representative of a new genus of Tethyidae. 


Materials and Methods 


Samples of Tethya (242 individuals) rep- 
resent part of several collections obtained 
along the Pacific coast of Mexico, including 
the Gulf of California at different depths 
(Table 1, Fig. 1). Sponges were collected 
with SCUBA diving and with a trawl 
aboard the R/V El Puma. Thirty-eight 
trawls, covering almost all the Gulf of Cal- 
ifornia from 31°15’9"N, 114°21'7’W to 
20°49'N, 105°42’W, from 22 to 120 m 
depth, were performed in March, July, Au- 
gust, and October of 1985. Several dives off 
the coast of Mazatlan took place from 
23°15'N, 106°29’W to 23°11'30"N, 106° 
25'W in May 1981 and June 1987 at a 
depth of 0 to 15 m. Trawls and dives from 
Guerrero were performed in February 
(Winter) and April (Spring) of 1982, in the 
continental shelf, from the coastline to the 
beginning of the continental slope, at 200 
m depth (102°15'’N, 98°W), including the 
Balsas river delta. Sponges, once aboard, 
were preserved first in 95% alcohol and 
then in 70%. Techniques to analyze internal 
structures follow Sara (1992), and scanning 
electron microscopy (SEM) follow Gomez 


VOLUME 114, NUMBER 3 


(1998). Spicular data reported are based on 
50 measurements from each spicular trait. 
Main and auxiliary megascleres, generally, 
strongyloxeas, were distinguished by a con- 
ventional 1000 pm length figure. For two 
species, a third category of “‘sword”’ stron- 
gyloxeas was considered. Micraster nomen- 
clature is traditionally based on observa- 
tions taken at the light microscopic level. 
Yet, the difference between oxyasters, 
strongylasters and tylasters (Sara 1994) is 
only partially due to the ray shape. It de- 
pends largely, as shown by SEM micro- 
graphs, on the distribution and strength of 
the spines. The holotypes are deposited in 
the Natural History Museum of Genoa (It- 
aly) (MSNG), some paratypes are deposited 
in Laboratorio de Ecologia de Bentos 
ICML collection of Mazatlan (Mexico). 
Study area.—The Gulf of California be- 
longs to the subtropical regime with marked 
fluctuations in climatic conditions all year 
long. It is positioned between two dry con- 
tinental environments that cause wide rang- 
es in temperature, low humidity, and high 
evaporation rate. The yearly mean temper- 
ature value at the surrounding coast is 24°— 
26°C. Rainfall is abundant at the east coast 
of the Gulf as well as in the south, whereas 
at the west coast rainfall is lower. Maxi- 
mum rainfall is from June to October, be- 
coming dry at winter and at the beginning 
of spring. This brings about coastal up- 
welling along the western side in summer 
and at the eastern side in winter (INEGI 
1984, Molina-Cruz 1986). Mazatlan is lo- 
cated at the mouth of the Gulf of California, 
in a tropical and subtropical semi-humid 
climate, bathed by the Gulf of California 
current and causing different water changes 
according to the nearby inlets, islands, and 
breakwaters. Yearly mean temperature is 
from 25°C to 28°C, yearly mean rainfall 
reaches 850 mm (Alvarez-Leon 1980). 
Guerrero has a tropical semi-humid climate, 
yearly mean temperature of 27.5°C, yearly 
mean rainfall 1117 mm, and a permanent 
influx of freshwater in Petacalco Bay that 


Table 1.—Sampling stations of Tethya species along the Pacific coast of Mexico. Collecting parameters. 


Guerrero 


Mazatlan 


Gulf of California 


Petacalco Bay & 


Punta Arboleda Punta Chile 


Cabo San Miguel 


Puerto Libertad 


Locality 


Pta. Maldonado 
17°55’N, 102°01’W 


29°59'0"N 28°10'2"N 26°51'9"N 23°12'28"N 


Coordinates 


16°13’N, 98°44'W 


45 both 


106°25’50”"W 


5&8 


110°00'9"W 


DD) 


112°48'2"W 


25 


112°45'0"W 
Intertidal 


Depth (m) 


Feb & Apr 1982 
21.7 & 23.5 


2M 
34.2 


May 1981 & Jun 1987 


22 


Jul 1985 


29.5 
4.8 


Mar 1985 


Oct 1985 


Collecting date 


14 
4.0 


17 
6.16 


Temperature (CC) 
Oxygen (ml/l) 


3.5 


35.5) 34.4 


35.56 35.3 


Sand 


Salinity (%o) 


Bottom 


Sandy-mud, clay 


Rock 


Irregular 


Rock-sand 


795, 


796 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
iL 115% 110° 105° 100° 95° © 9p 
‘ : ro te 
. 
| S 
—— \ P. Libertad 0 
SS _ ® T socius : ea 
ON we - waa 
ee Miguely 6) ~~ a 
= calt ee e ig a 
7 » P. Arboleda og! 
ie. 2 pOew Mm. T paroxeata q 
bo Q. ¢ IL mexicana 5 
SR 2 
& i 4 % %s Se M e = & 
“I oo ( : 
te. €XICO i Gulf of Mexico ™ 
4 “\ Mazatlan 
Vv TF taboga 
\ q 
a ( ‘eee 
= : f 
° be ne 5 ° 
R East Pacific 7. oN ; 
J = JR. Balsas i. eee, 
ee ie ae 
B. Petacalco a 
y mexicana ER oe 
- ovum e hae 
P. Makionadd® es 
e T. mexicana SS ° 
= ° 5 T. ensis “Se nee 
HS Or 105° | (OOR 7 ess Se 90° 
Fig. 1. Map of collecting areas and distribution of Tethya species: Puerto Libertad, Cabo San Miguel and 


Punta Arboleda inside the Gulf of California, Mazatlan at the mouth, and Petacalco Bay as well as Punta 
Maldonado at Guerrero, southern Mexico, tropical East Pacific. 


comes from the Balsas river (Amezcua-Lin- 
ares 1996). 

The study area, except the Gulf of Cali- 
fornia, represents the transition zone be- 
tween the mixture from the North Califor- 
nian current and the South Peru current 
with the North equatorial counter-current, it 
marks the limits between the Californian 
Province with the Panamic one, now sub- 
divided in Cortez Province, Mexican Prov- 
ince, and Panamic Province (Hendrickx 
1995). 


Systematics 


Order Hadromerida Topsent, 1900 
Family Tethyidae Gray, 1867 
Genus Tethya Lamarck, 1815 

Type species.—Tethya aurantium Pallas, 
1766. 

Diagnosis.—Spherical or subspherical 
body not supported by a stalk, well devel- 


oped cortex distinct from the choanosome, 
megascleres bundles radiate from the center 
of the sponge ending in tubercles on the 
surface. Main and auxiliary megascleres are 
usually strongyloxeas, auxiliary may be 
also styles, megasters are spherasters or ox- 
yspherasters, micrasters are tylasters, stron- 
gylasters (chiasters) or oxyasters; these two 
are distributed in the cortex as well as in 
the choanosome (Sara 1994). 


Tethya ensis, new species 
Figs. 2A, B, 3, 4, 15A, Table 2 


Material examined.—Holotype: MSNG 
50191; paratype: LEB-ICML 255; 46 spec- 
imens Punta Maldonado (Guerrero), Feb 
1982, 45 m depth. 

Description.—The type has an elongated 
hemispherical shape, 1.5 by 2.5 cm, with 
the irregular flattened basis covered by ro- 
bust filamentous stolons. On the whole, the 


VOLUME 114, NUMBER 3 


shape is elongated hemispherical (Fig. 2A, 
B), 0.8—2.6 by 0.7—3 cm. Two specimens 
were attached to the next Tethya species de- 
scribed herein. Color: not recorded in vivo, 
creamy white in ethanol. Consistency 
slightly compressible. Surface sparsely and 
irregularly tuberculate, partially smooth. 
The best distinguished tubercles are finely 
hispid, flattened, 1—2 mm in diameter, 1 mm 
in height (Fig. 2A), sharp and stout fila- 
mentous stolons on the basis and the edges 
of the sponges are 2 cm long. Some tuber- 
cles on the edge of the sponges produce 
also 3 mm elongated buds. Cortex thick- 
ness, without tubercles, 0.5—1 mm. 

Skeleton.—Megasclere bundles radiate 
from center to cortex sometimes in a coiled 
way as in the holotype. Bundles, 245—325 
zm in diameter, ending in compact cortical 
fans without subdivisions (Fig. 3A). Me- 
gasters regularly distributed in the middle 
and, more densely, in the lower cortex, 
forming a belt around the choanosome. 
Some smaller megasters in the outer choan- 
osome (Fig. 15A). 

Spicules.—Table 2 summarizes measure- 
ments taken from 5 specimens. Megascleres 
(Figs. 3B, D, 4A) are: main strongyloxeas 
(maximum size 2312 by 47 wm), peculiar 
shortened fusiform strongyloxeas (“‘sword”’ 
like) in the cortical fans and among the sub- 
cortical interstitial megascleres, 225—965 
jum, with a slender head 4—8 ym thick and 
a thickness of generally 15—25 «wm, in the 
central and distal parts of the spicule and 
auxiliary megascleres, from slender stron- 
gyloxeas to thin styles. 

Megasters (Figs. 3D, 4D): Spherasters 
heterogeneous in size and shape, with some 
slight differences among the specimens and 
a main size range 50—90 wm (maximum di- 
ameter 115 pm). The R/C main range is 
0.5—0.9, sometimes in the larger spicules it 
is 1—-1.1. Ray number: 14-16. 

Micrasters (Figs. 3C, 4B, C): Mainly 
strongylasters but variable from slightly 
knobbed tylasters to slightly tylote oxyas- 
ters 9-13 wm in diameter (minimum 5.9 
and maximum 16.5 wm) with 12 thin spiny 


TST 


rays. Similar in the cortex and in the choan- 
osome. 

Etymology.—From Latin ensis = sword, 
in reference to the sword-like appearance of 
the short fusiform strongyloxeas; here used 
as a noun in apposition. 

Remarks.—Tethya ensis is characterized 
by a peculiar shape and the occurrence in 
the cortex of the sword shortened fusiform 
strongyloxeas. The species is akin to a 
group of Mexican species subsequently de- 
scribed herein and to 7. californiana De 
Laubenfels, 1932 (Sara & Corriero 1993). 
Tethya ensis differs from T. californiana in 
body shape and cortical structure and in 
some significant spicular traits: the mega- 
scleres are strongyloxeas instead of aniso- 
strongyles (but a Mexican population of 7. 
californiana subsequently described herein 
also has strongyloxeas); the occurrence of 
peculiar sword shortened strongyloxeas 
which are absent in 7. californiana; and the 
larger size and the lower R/C of its megas- 
ters that are spherasters instead of oxy- 
spherasters. 


Tethya mexicana, new species 
Figs. 2C—G, 5-7, 15C, Table 3 


Material examined.—Holotype: MSNG 
50192; paratype: LEB-ICML 256; 75 spec- 
imens from Punta Maldonado (Guerrero), 
Feb 1982 and 10 specimens from Petacalco 
Bay (Guerrero), Apr 1982, 45 m depth. Ad- 
hered to bryozoans and calcareous frag- 
ments on sand and sand-mud-clay bottom; 
43 specimens from Punta Arboleda (Gulf of 
California), Jul 1985, 22 m depth. 

Description.—The type is irregularly el- 
lipsoidal, cushion-like, with two opposite 
faces 3 by 2 cm, both covered by irregularly 
rounded tubercles, 1-3 mm in diameter, 
0.5—2 mm in height. The thickness of the 
cushion is 1.5 cm. Long filamentous rooting 
processes are at one side of the faces, sug- 
gesting that in life the specimen was erect 
2 cm high and 3 cm broad. Some elongated 
tubercles with spear like buds are on the 
top. On the whole, body variable in shape, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2._ A, B: Tethya ensis, new species. A, Holotype; B, Paratype. C-G: Tethya mexicana, new species. C— 
E, Three specimens from Gulf of California; E Holotype from Guerrero; G, Transverse section of another 
specimen from Guerrero showing coiled skeleton and a central nucleus. H—J: Tethya paroxeata, new species. 
H, Holotype lateral view; I, Holotype, transverse section showing a coiled skeleton radiating from an asym- 
metrical center; J, Another specimen, lateral view. Scale bars = 1 cm. 


VOLUME 114, NUMBER 3 


Fig. 3. Tethya ensis, new species. Light microscopic photographs. A, Skeletal structure of tubercle; B, 
Fusiform strongyloxeas and some megasters; C, Micrasters; D, Megasters and megasclere endings. Scale bars 
= 100 pm. 


800 


Table 2.—Measurements of spicules of specimens of Tethya ensis (wm). Underlined numbers indicate mean values. 


Megascleres 


Auxiliary Megasters 


“Sword” strongyloxeas 


Main 


Micrasters 


Diameter 


Diameter R/C 


Length Width 


Length Width 


Length Width 


Specimens 


5.9-10.5-16.5 
8.3-11.6-15.3 


0.7-1.0 


0.4-0.6-0.8 


0.4 


37-81-106 
27-87-115 


3-12-28 
3-9-22 


381-571-972 
318-603-980 


12-1726 
t= 


319-443-609 
364-510-720 


15-30-47 
8-24-45 


1056-1611-2115 
1006-1623-2312 


1 (Holotype) 


2 


9.0-12.1-14 


va 0.5-0.7-1.1 


42-77-106 


4-8-20 


300-545-970 


10-15-22 


270-493-780 


6-26-36 


1005-1586-2280 


10.0-11.9-13.5 


0.4-0.6-0.9 


46-76-98 


4-7-13 


260-551-990 


10-16-21 


247-513-777 


18-29-35 


1342-1714-2290 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


4-0.6- 


34-67-106 


4-10-23 


390-735-985 


10-15-23 


225-472-965 


10-27-36 


1000-1599-2100 


0. 


with irregularly hemispherical, ellipsoidal 
or polyhedral specimens, 0.8—4 cm in di- 
ameter and 0.5—2.5 cm in height (Fig. 2C— 
G). Color orange when alive, dirty white or 
brownish in ethanol. Consistency firm, 
slightly compressible. Surface tuberculate, 
with irregularly rounded or roughly polyg- 
onal tubercles 1—3 mm in diameter, 0.5—2 
mm in height irregularly spaced and un- 
evenly developed in Guerrero specimens, 
sometimes flattened, sometimes papillose, 
covered by sand in the Gulf of California 
specimens, others are scarcely visible with 
a surface covered by several stalked buds 2 
cm long, and several with visible oscules. 
Specimens from. Guerrero, as in the type, 
have on the basis and edges of the body 
flattened or filamentous rooting processes 
10—16 mm long by 1-10 mm wide. Cortex 
(Fig. 5C, D), including tubercles, 2—4 mm 
thick. Spicular tufts hispidating the tuber- 
cles supported by bundles of strongyloxeas 
with a diameter of 300—480 pm in the cor- 
tex and 350—950 p.m in the choanosome. 

Skeleton.—The megasclere tracts branch 
at different levels of the choanosome (Fig. 
5B). The interstices among the tracts in the 
upper part of the choanosome are filled with 
bundles of auxiliary megascleres. Megas- 
ters are placed densely in the central and 
lower parts of the cortex that shows several 
lacunes (Fig. 15C), although it is less la- 
cunar than Tethya californiana. In some 
specimens the megasclere tracts are coiled 
and sometimes they depart from a central 
nucleus (Fig. 2G). The nucleus, about 5 mm 
in diameter, is made up of an irregular net- 
work of small styles and subtylostyles (Fig. 
SA). 

Spicules.—Table 3 summarizes measure- 
ments taken from three specimens of Guer- 
rero and three specimens from the Gulf of 
California. Megascleres (Figs. 6A, 7B) are 
main strongyloxeas and auxiliary (cortical) 
strongyloxeas with intermediates among the 
two categories. Maximal length is 2130 wm 
and thickness 41 wm. The nucleus of some 
specimens presents heterogeneous styles 


VOLUME 114, NUMBER 3 


801 


Fig. 4. Tethya ensis, new species. SEM micrographs. A, Megascleres and some megasters; B, Tylaster (more 
frequent); C, Chiaster (less frequent); D, Spheraster. 


and subtylostyles frequently somewhat 
curved, 300—800 by 10-18 pm. 

Megasters (Figs. 6A, 7C) are spherasters- 
oxyspherasters: heterogeneous in size and 
shape, they vary also among the species. 
The main size range is 70-120 wm (maxi- 
mum 128 wm) and R/C range 0.7-1.2 
(maximum 1.4). Two specimens from Punta 


Arboleda (Gulf of California) show sharp 
differences. The main size range is 60—75 
wm and the main R/C range respectively, 
0.5—0.7 and 0.7—0.9. Ray number 16-18, 
frequently bent, sometimes twisted or 
forked. 

Micrasters: Mainly strongylasters (chias- 
ters) but variable from slender tylasters with 


802 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 5. Tethya mexicana, new species. Details of structure. A, Nucleus skeleton; B, Branching of a mega- 
sclere bundle in the choanosome; C, Cortical structure; D, Cortical structure with megasclere fans. Scale bars 
= 100 pm. 


VOLUME 114, NUMBER 3 


Table 3.—Measurements of spicules of specimens of Tethya mexicana (wm). Underlined numbers indicate mean values. 


Megascleres 


Megasters 


Auxiliary 


Main 


Micrasters 


Diameter 


Diameter R/C 


Length Width 


Width 


Length 


Specimens 


1 (Holotype) 


6.8-11.4-15.5 


8-1.2 


339-547-971 4-14-26 48-97-127 


14-23-40 


1020-1339-2010 


(Guerrero) 
2 (Guerrero) 
3 (Guerrero) 


8.0-11.6-15.5 


36-85-108 


5-16-38 


359-670-990 


14-26-36 


1000-1522-2070 


0.7-1 


8.5-11.6-15.5 


70-102-130 


5-15-23 


360-656-985 


18-29-39 


1033-1428-1820 


8.0-11.3-15.0 


49-64-78 


4-12-23 


347-745-990 


16-29-41 


1000-1564-2105 


4 (P. Arboleda) 
5 (P. Arboleda) 
6 (P. Arboleda) 


8.0-11.3-15.0 


23-59-93 


4-12-21 


250-725-978 


9-28-39 


1000-1568-2130 


5.2-8.7-12.5 


59-99-128 


6-15-24 


494-808-999 


10-20-34 


1002-1340-2021 


0.6-0.9-1.3 


803 


slightly knobbed rays to slightly tylote ox- 
yasters. Sometimes with a more or less de- 
veloped center. They are in the cortex and 
in the choanosome alike, generally 8—13 
jum in diameter (maximum 15.5 and mini- 
mum 5.2), with 8-14, generally 12, thin 
rays (Figs. 6B, 7A, D). 

Etymology.—Named after its origin, the 
Mexican waters. 

Remarks.—Tethya mexicana is charac- 
terized by a variable, generally depressed, 
body shape, uneven tuberculate cortex, 
sometimes with long stolons as in 7. ensis, 
sometimes with buds, large and variable in 
size, spherasters-oxyspherasters with rays 
frequently bent, micrasters similar in the 
cortex and in the choanosome varying from 
a Slightly tylote oxyaster to strongylaster 
shape or true slender tylasters. There is 
some intraspecific variability in spiculation, 
with differences especially among the spec- 
imens of Guerrero and Punta Arboleda. One 
of the Arboleda specimens (6) is character- 
ized by smaller megascleres and micrasters 
(Table 3). The general structure and the 
spicular traits have several affinities with 7. 
ensis but also some remarkable differences: 
the body shape even if variable, is not elon- 
gated hemispherical as in T. ensis; the skel- 
etal structure shows a branching of megas- 
clere tracts and interstitial strongyloxeas 
which are absent in 7. ensis; the sword-like 
strongyloxeas of T. ensis are lacking in T. 
mexicana; the megasters (spherasters-ox- 
yspherasters) are considerably larger in T. 
mexicana and with a greater R/C than in T. 
ensis. Tethya mexicana is also similar to T. 
californiana but differs in some important 
traits: the body shape is not spherical, has 
uneven tubercles and different cortical 
structure; the denser distribution of megas- 
ters in the lower cortex; the presence of 
strongyloxea instead of anisostrongyle type 
of megasclere; a greater size, lower R/C and 
lower ray number of its megasters; the more 
slightly tylote oxyaster type of its micras- 
ters; and the lack of spherules. 


804 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. 
bars = 100 pm. 


Tethya ovum, new species 
Figs. 8A, B, 9, 10, 15B, Table 4 


Material examined.—Holotype MSNG 
50193; paratype: LEB-ICML 257; 4 speci- 
mens from Petacalco Bay (Guerrero), Apr 
1982, 45 m depth. 

Description.—The type is ovoidal 11 
mm in diameter and 17 mm in height. On 
the whole, 10-17 mm in diameter, 15—25 
mm in height (Fig. 8A). Color, not recorded 
alive, dirty white in ethanol. Consistency 
hard. Surface strongly tuberculate, tubercles 
irregularly spaced, uneven, microhispid, 
sometimes smooth: 1—4 mm in diameter, 1— 
2 mm in height. Cortex, including tubercles, 
3—4 mm thick. 

Skeleton.—Megasclere bundles less than 
500 wm in diameter, sometimes a little 


Tethya mexicana, new species. Light microscopic photographs. A, Megasters; B, Micrasters. Scale 


coiled, radiating from the center to the cor- 
tex (Fig. 8B). Smaller fusiform subtylote 
strongyloxeas occur in the tubercle fans. 
Megasters are placed mainly in the lower 
cortex as a narrow continuous belt around 
the choanosome (Fig. 15B). Micrasters are 
similar in the cortex and in the choano- 
some. 

Spicules.—Table 4 summarizes measure- 
ments taken from two specimens. Main and 
auxiliary megascleres are slightly subtylote 
strongyloxeas. Maximum size of main me- 
gascleres is 1885 by 39 ym. Shortened cor- 
tical sword-like megascleres are fusiform, 
with the head 8—10 pm thick and the great- 
er thickness 10—27 pm, in the central and 
distal parts of the spicule (Figs. 9A, 10A). 

Megasters: Spherasters and oxyspheras- 


VOLUME 114, NUMBER 3 


X53, 666 


1g@kU %1,006 


Fig. 7. 


805 


Tethya mexicana, new species. SEM micrographs. A, D. Oxyasters with slightly tylote ends (more 


frequent); B, Megascleres and several megasters; C, Megaster. 


ters show a main size range of 75-100 wm 
(maximum 110 wm), and R/C 0.7-1.2 
(maximum 1.5), with about 18 rays gener- 
ally bent and frequently spined or bifid 
(Figs. 9B, 10D). 

Micrasters: Mainly strongylasters (chias- 
ters), but they vary from slender tylasters 
with slightly knobbed rays to slightly tylote 


oxyasters. Size: 8-13 wm in diameter, with 
about 12 thin rays, in the cortex and choan- 
osome alike (Figs. 9C, 10B, C). 
Etymology.—From latin ovum = egg, in 
reference to the ovoid shape of the body, 
an unusual character in Tethya; here used 
aS a noun in apposition. 
Remarks.—Tethya ovum is similar to T. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 8. A, B: Tethya ovum, new species. A, Lateral view of paratype; B, Transverse section of the same. C, 
D: Tethya socius, new species. C, Upper view of holotype; D, Basal view of the same. E, F: Tethya californiana 
De Laubenfels, two specimens showing differences of tuberculate surface. G: Tethya taboga (De Laubenfels) 
upper view. Scale bars = 1 cm. 


VOLUME 114, NUMBER 3 807 


ensis and to T. mexicana but is well char- 
acterized by its ovoidal shape and strongly 


S 

Ballas tuberculated surface. It shares with T. ensis 
EE eI the sword-like category of strongyloxeas, 
ar) an but these are clearly distinct because their 
‘© 26 head diameter is 8-10 ym instead of 4-8 
wm. The megasters are spherasters-oxys- 

NM . O : 
= pherasters similar to those of 7. mexicana 
S ale | but generally bent and frequently adorned 
we by forks and spines while the skeleton 


structure is more alike to T. ensis. Tethya 
ovum differs from T. californiana in all the 


Megasters 


Yo) &) 
: ae traits discussed for 7. ensis and T. mexi- 
a = ~ ISI cana. 
3 as 
g 
g Tethya paroxeata, new species 
= S| S« Figs. 2H—-J, 11, 12, 15D, Table 5 
io e | Wile 
8 pace Material examined.—Holotype MSNG 
iE Ey e 50194; paratype LEB-ICML 258; 6 speci- 
iS a) 
x E % A 36 mens from Punta Arboleda (Gulf of Cali- 
E #! S\s| fornia), Jul 1985, 22 m depth. 
e || ose Description.—The type is irregularly 
g Sa polyhedral 2 cm high with flattened basis 
3 and six faces, 2 by 2 cm. The origin of the 
5 | aq radiate bundles of the megascleres is a cen- 
zB 4 3 <2 | tral nucleus near the surface of a lateral 
& g =F face, and the bundles are then strongly 
Saale, 2 coiled towards the opposite face (Fig. 21). 
e 13)? 22 In other four specimens, the body shape is 
S| E al dia irregularly polyhedral 1.5—3 cm at the base, 
& [712 |s ele 12.5 cm in height (Fig. 2H,J), and in two 
i) to a specimens hemispherical with a diameter of 
a 2.5 cm. Color orange when alive. Consis- 
a ee tency compact but not hard. Surface with 
2. e| o S| small flattened and contiguous tubercles, 1— 
Bileay a 1.5 mm in diameter in two specimens, larg- 
o = er (2 mm) but more flattened and surface 
8 2 ire nearly smooth in the other three specimens. 
fa 5 2 gs Many sand particles on the surface. With 
a Fp ees stalked buds (stalks 1 cm long). Tubercles 
3 5 ale 0.5 mm high, cortex (without tubercles) 1.5 
® ee mm thick, very lacunar under the sponge 
g == surface (Fig. 15D). 
= Skeleton.—Megasclere tracts, 280—630 
| p 2 yum thick, run compact to the tubercles 
2 || forming little-developed fans (Figs. 11A, 
S Bt = 15D). Some tracts, however, branch in the 
= inner part of the choanosome. Auxiliary 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 9. Tethya ovum, new species. Light microscopic photographs. A, Fusiform strongyloxeas and three 
megasters; B, Spined and forked megasters with parts of megascleres; C, Micrasters and points of megascleres. 
Scale bars = 100 pm. 


VOLUME 114, NUMBER 3 


809 


Fig. 10. Tethya ovum, new species. SEM micrographs. A, Megascleres and megasters; B, Tylasters (more 
frequent) with one chiaster at middle left; C, Chiaster (less frequent); D, Megaster. 


megasclere bundles occur in the upper 
choanosome in some interstices among the 
main megasclere tracts. Megasters are irreg- 
ularly and not densely distributed in the 
middle and inner cortex, more dense in the 
middle part (Fig. 15D). The surface is coat- 
ed with a dense layer of micrasters. 


Spicules.—Table 5 summarizes measure- 
ments taken from five specimens. Megas- 
cleres are fusiform strongyloxeas. Several 
middle-sized (1000-1300 by 10—20 pum), 
and the great majority of the small (300— 
1000 by 1—20 pm) megascleres are oxei- 
form with a very thin (0.5—3.5 jm thick) 


810 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


rounded or tylote head (Figs. 11C, 12B). It 
is not easy to distinguish between main and 


auxiliary megascleres because there is a aco ee 
continuous range in their length (283—2620 28 =a ae 
um), thickness (1-50 jm) and head width b2| Q/2(e|=I-| 
(0.7—22 wm), and these parameters are not mOCooSe 

wt wt CO 00 © 


always correlated. Moreover, large and 
small megascleres are associated in the 
same bundle. Some measurements (head 
thickness in brackets): 2620 by 45 wm (15), SSS = 


1 
0) 
0) 
3 
5 


2500 by 45 um (18), 2250 by 50 jum (20); 2| 4lslsléle 
1880 by 40 wm (10); 1840 by 40 wm (10); pe 
1550 by 38 pm (22); 1240 by 10 pm (2), é : 
950 by 20 pm (5); 900 by 7 wm (1.5); 830 S 2 
by 8 pm (2.5); 770 by 6 pm (1); 670 by 5 s 2 ae 
uum (1). There are also strongyles, (one with S : 3 Ws aie 2 
the rounded head of the transformed stron- 2 a tanta eS 
gyloxea a little tylote, 1050 by 35 jm (20), =| oh 
another 1135 by 35 wm (15) and anisotron- 3 
gyles. The measurements reported in Table 2 
5 refer to a conventional figure to distin- : «| o228n 
guish between main and auxiliary megas- se) : Ot [oh [cu 
cleres at a length of 1000 pm. The larger = aAOoONnW 
megascleres (maximum size 2620 by 50 3 & 
xm) have a head generally slightly tylote =) s 
thick 10—22 jm, and a distal end generally -) Z LI SHS 
well pointed, sometimes slightly rounded. = a ear, 
In contrast, the middle-sized and smaller iS g ZIEBIRIS| 
megascleres are also fusiform, but looking S = = a =) 5 
like oxeas for their very thin (0.5—3.5 pm) & 
and generally slightly tylote head. & [8 
Megasters (Figs. 11D, 12A): Spherasters, a 
sometimes oxyspherasters, with the main SS . Bat Fm 
range 40-70 pm (maximum 91 pm) in di- a 2 alalalslal 
ameter. R/C range, more frequently 0.6—1 = See 
(maximum 1.5). Ray number: 16—20, rays 2, 
sometimes slightly bent or bifid. Spheras- ‘es ¥ 
ters, not abundant, are variable in size and 6 3 EMS SS 
shape in the same specimen and among the 8 6 & a A = 
specimens. 2 2] Siz Vlaje 
Micrasters (Figs. 11B, 12C, D): Similar 5 8 Sle lelale 
in the cortex and in the choanosome. Main- 3 = = S =) =) 
ly strongylasters (chiasters) but variable ©) Inblalolo” 2a 
from tylasters with slightly knobbed rays to a 
slightly tylote oxyasters. Size: 8-13 pm = 
(minimum 4, maximum 15.5 wm) in di- il : 2 
ameter. Ray number: 12—14. Often a center 2 Pa ee 
is more or less developed. s Z = 
Etymology.—From the Greek prefix para a aAM tH 


VOLUME 114, NUMBER 3 


Fig. 11. 
C, Slender strongyloxeas; D, Megasters. Scale bars = 100 pm. 


= beside, nearby + oxeata = relating to an 
oxea, Neolatin from Greek oxys = sharp, in 
reference to the ends, both thin, of the small 
megascleres. 

Remarks.—Tethya paroxeata is similar 
to T. ensis, T. mexicana and T. ovum but 
is clearly distinguished by its peculiar 
polyhedral shape and even surface. The 
same type of shape and surface may occur 
rarely in some specimens of T. mexicana 
from the same locality (Punta Arboleda) 
but less pronounced. Tethya paroxeata is 
clearly distinguished from similar species 


Tethya paroxeata, new species. Light microscopic photographs. A, Cortical structure; B, Micrasters; 


by the peculiar oxeiform shape of the 
greater majority of the shorter megascler- 
es. The spherasters, rarely oxyspherasters, 
are smaller than in 7. ensis, T. mexicana 
and 7. ovum. Spicular intraspecific vari- 
ability is shown by specimen two which 
is characterized by smaller micrasters (Ta- 
ble 5). Tethya paroxeata differs from T. 
californiana in body shape, cortical sur- 
face and structure and occurrence of the 
oxeiform megascleres, in addition to the 
other traits listed for T. ensis, T. mexicana 
and 7. ovum. 


812 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 12. Tethya paroxeata, new species. SEM micrographs. A, Megaster; B, Megascleres; C, Strongylasters 
(less frequent); D, Tylaster (more frequent). 


Tethya socius, new species 
Figs. 8C, D, 13, 14, 15E, Table 6 


Material examined.—Holotype MSNG 
50195; 1 specimen (composed of 4 fused 
individuals) from Puerto Libertad (Gulf of 
California), Oct 1985, intertidal. 

Description.—The type is made by 4 
fused spherical depressed individuals of 2, 


1.5, 1.2 and 1 cm in diameter and 1—1.2 cm 
in thickness. On the whole, the specimen 
measures 4 by 3 by 1—1.2 cm in thickness 
(Fig. 8C, D). Color: pale pink in ethanol. 
Consistency soft, rather compressible. La- 
cunes in the outer cortex and in the outer 
choanosome under the cortex (Fig. 15E). 
Surface with contiguous tubercles of irreg- 


VOLUME 114, NUMBER 3 


ular shape and different size, 1-3 mm wide, 
flattened and with slightly winding outlines. 
Cortex with the tubercles, 2—2.5 mm thick. 
The body of all the individuals has a strong 
central nucleus, about 6 mm in diameter, 
from which the megasclere bundles radiate. 

Skeleton.—Yhe megasclere tracts may 
split, as in 7. mexicana, before reaching the 
cortex. Different megasclere bundles may 
also support the same tubercle. Terminal 
fans are little developed. Auxiliary megas- 
clere bundles fill the interstices among the 
main tracts in the upper choanosome. 
Spherasters are distributed in all the cortex, 
more densely in the basal part, especially at 
the boundary with the choanosome. Some 
smaller megasters are in the outer part of 
the choanosome. The skeleton of the nucle- 
us is made by a dense and irregular network 
of small styles and subtylostyles (Fig. 15E). 

Spicules.—TYable 6 summarizes measure- 
ments taken from the holotype. Megascler- 
es (Fig. 14A): strongyloxeas little fusiform 
without a clear distinction between the 
main and auxiliary. In the table the two cat- 
egories are conventionally separated on the 
basis of a 1000 wm length figure. Some 
measurements (head thickness in brackets): 
1700 by 30 wm (10); 1350 by 18 pm (10); 
1315 by 18 wm (10); 1250 by 22 pm (12); 
1040 by 8 pm (5); 1030 by 18 wm (5); 700 
by 2 wm (1). An anisostrongyle: 1460 by 
— 28 pm (15; 10). Maximal sizes 1700 pm in 
length and 30 wm in thickness, on the 
whole: 440-1700 pm by 2-30 pm (1-12). 
In the center small styles or subtylostyles 
heterogeneous in length and thickness and 
sometimes with the basal third a little 
curved: 150—400 by 5—20 pm. 

Megasters (Figs. 13B, C, 14B, D): Ox- 
yspherasters: generally 60-100 (max. 118 
um in diameter); R/C generally 1.2—1.7 
(max. 1.8, min. 0.7). Ray number: about 14, 
frequently bent, twisted, blunt, bifid or with 
spines on the rays. 

Micrasters (Figs. 13B, 14C): Mainly 
strongylasters but variable from slightly 
knobbed tylasters to slightly tylote oxyas- 
ters. They are similar in the cortex and in 


Table 6.—Measurements of spicules of one specimen of Tethya socius (4m). Underlined numbers indicate mean values. 


Megascleres 


Megasters 


Auxiliary 


Main 


Micrasters 


Diameter 


Diameter R/C 


Length Width 


Width 


Length 


Specimens 


0.7-1.3-1.8 6.5-10.1-14.6 


DAME) 20-71-118 


8-18-30 439-768-992 


1003-1206-1700 


1 (Holotype) 


813 


814 


Fig. 13. 
one megaster; C, Megasters. Scale bars = 100 pm. 


the choanosome, generally 8—12 wm in di- 
ameter. Ray number 10-14. 
Etymology.—From Latin socius = com- 
panion, in reference to the fusion of differ- 
ent individuals to form one specimen, un- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Tethya socius, new species. Light microscopic photographs. A, Cortical structure; B, Micrasters and 


usual in Tethya; here used as a noun in ap- 
position. 

Remarks.—Tethya socius is well distin- 
guished from the other Mexican species 
here described for the smaller size of the 


VOLUME 114, NUMBER 3 


p 


ae 


Fig. 14. Tethya socius, new species. SEM micrographs. A, Megascleres; B, Megaster; C, Micraster (more 


frequent); D, Megaster. 


megascleres, the very high R/C of its ox- 
yspherasters with rays frequently twisted, 
bent, blunt, bifid or with spines. Only the 
micrasters, similar in the cortex and in the 
choanosome as in the other Mexican spe- 
cies, are roughly alike to those of T. ensis, 
T. mexicana, T. ovum, and T. paroxeata. 
The frequent oxyasters with a small swell- 
ing at the ray tips, which occur in all these 


Mexican species and also in 7. californi- 
ana, may be compared to a similar trait in 
the eastern Indian Ocean T. affinis Kirkpa- 
trick, 1900 and to the western Pacific T. de- 
formis Thiele, 1905. Yet, in T. affinis the 
rays are longer and slenderer, and in T. de- 
formis the micrasters have a well developed 
center. Other spicular traits of both these 
species are clearly different by the Mexican 


816 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 15. Reconstruction of skeletal patterns from transverse sections of the new species of Tethya. A, T. 
ensis; B, T. ovum; C, T. mexicana; D, T. paroxeata; E, T. socius. 


species. Tethya socius is also characterized Borges, 1991 from S.E. Australia. But T. 
by the fusion of the subspherical individu- communis lacks the central nucleus and has 
als which show a well marked central nu- oxyspherasters with shorter rays and polyr- 
cleus. A similar fusion of individuals is habds, absent in 7. socius, among the mi- 
found in T. communis Bergquist & Kelly- croscleres. A central nucleus has been 


VOLUME 114, NUMBER 3 


found also in some specimens of 7. mexi- 
cana. 


Tethya californiana De Laubenfels, 1932 
Fig. 8E, E Table 7 


Tethya aurantia var. californiana De Lau- 
benfels, 1932:44 

Tethya californiana.—Sara & Corriero, 
1993:204 


Material examined.—54 specimens from 
Cabo San Miguel (Gulf of California), Mar 
1985, 25 m depth. 

Description.—Body shape spherical or 
sub-spherical, 0.6—3.8 cm in diameter. Sur- 
face smooth due to the very even tubercles. 
Color ochre yellow to orange when alive. 
Some specimens have round to irregular os- 
cules on top. Consistency soft in ethanol. 
Cortex including the tubercles 2—2.5 mm 
thick. Tubercles 0.5—-1.5 high, 1-3 mm 
broad. The irregular outlines of tubercles 
and the lacunar structure of the cortex and 
the choanosome correspond to those in the 
redescription of 7. californiana in Sara & 
Corriero (1993). 

Megasclere bundles 300—800 wm in di- 
ameter. 

Spicules.—Measurements of 5 specimens 
are summarized in Table 7. Megascleres are 
main and auxiliary strongyloxeas with in- 
termediates between the two categories. 
' Maximal size of strongyloxeas is 2375 by 
43 wm. Medium-sized and small strongy- 
loxeas have often the proximal part (head 
and neck) a little bent on the axis of the 
spicule. 

Megasters: are mostly spherasters with a 
wide range in size and in R/C, even in the 
Same specimen. Generally measuring be- 
tween 45 and 60 pm, but minimal and max- 
imal diameters are 20 and 83 wm. R/C gen- 
erally between 0.6 and 0.8 but minimal and 
maximal R/C are 0.3 and 1.2. The conical 
rays about 24, are sometimes bifid or blunt. 

Micrasters: similar in the cortex and in 
the choanosome are mainly microspined 
strongylasters, frequently as a strongylaster- 
oxyaster type, with moderately pointed 


Table 7.—Measurements of spicules of specimens of Tethya californiana (wm). Underlined numbers indicate mean values. 


Megascleres 


Megasters 


Auxiliary 


Main 


Spherule 
Diameter 


Micrasters 


Diameter 


Diameter R/C 


Width Length Width 


Length 


Specimens 


25-50-68 0.4-0.7-1.2 7.9-11.3-14.8 


1-4-18 


407-725-983 


12-21-36 


1053-1373-2017 


6.7-10.0-13.5 


0.4-0.7-1.1 


23-51-66 


17-22 


286-686-976 


7-26-43 


1010-1742-2375 


8.0-11.0-13.0 


8.0-10.2-13.0 
8.0-10.2-13.0 


8.0-11.8-15.5 


9.0-12.3-15.5 


7.5-11.7-15.0 


0.4-0.6-1.0 
0.4-0.7-1.1 


0.3-0.6-0.09 


26-54-83 
21-54-78 


20-55-75 


3-11-18 
4-13-28 


Ba WD) 


270-738-995 


307-698-990 


270-777-995 


10-26-40 
10-26-40 


10-29-40 


1000-1402-2090 


1000-1576-2340 


1000-1604-2100 


Norn 


817 


818 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


rays, and, less frequently, as a strongylaster- 
tylaster type, with slightly knobbed rays. 
Ray number 10—14 and, sometimes, a small 
center. More frequent size: 10—13 wm (max. 
15.5 and min. 6.7). 

Spherules 8-13 wm, are present in the 
cortex. 

Remarks.—The Mexican population was 
identified as Tethya californiana after com- 
parison with the type material of the spe- 
cies. However, it differs considerably in two 
traits: the megascleres are strongyloxeas in- 
stead of anisostrongyles and the megasters 
are generally spherasters, with R/C lower 
than 1, instead of oxyspherasters. Yet, some 
oxyspherasters with R/C = 1—1.2 occur also 
in the Mexican population. Other slight dif- 
ferences between the Mexican population 
and the Pacific 7. californiana are the 
smaller body size, the smoother surface, 
and the greater spherule diameter. These 
differences may be due to the different geo- 
graphic location of the samples. The type 
material comes from the Pacific coast of 
California, north of Los Angeles, whereas 
the ones described here are from the Mex- 
ican coast of the Gulf of California. 


Oxyasters 
Diameter 
11.5-45.6-67.5 
22.3-42.9-58.5 
22.2-39.2-55.0 


Micrasters 

Diameter 
6.6-9.5-13.4 
6.3-8.7-12.7 


8.5-10.4-12.5 


Megasters 
R/C 
0.3-0.5-0.8 


Diameter 
19-48-65 
42-62-74 


30-57-70 


Width 


4-14-30 
2-9-19 
1-12-25 


Tethya taboga (De Laubenfels, 1936) 
Fig. 8G, Table 8 


Taboga taboga De Laubenfels, 1936:452 
Tethya aurantia sensu Green & Gomez, 
1986:284 


Auxiliary 


Length 
182-698-980 
262-587-987 
323-600-987 


Material examined.—Punta Chile (Ma- 
zatlan), 7 specimens, May 1981, 8 m depth, 
2 specimens, Jun 1987, 5 m depth. 

Description.—Body shape hemispheri- 
cal, 2.5 cm in diameter, 1.5 cm in height. 
Surface with rounded tubercles 1—1.5 mm 
high and broad. Cortex including the tuber- 
cles, 2—2.5 mm thick. 

Megasclere bundles are sometimes coiled 
from the center to the cortical surface. 

Spicules.—Measurements of three speci- 
mens are summarized in Table 8. Megas- 
cleres are generally strongyloxeas, some- 
times with stepped ends. Maximal size 
found for strongyloxeas is 1680 by 34 wm. 


Megascleres 


Width 
16-27-34 
12-18-28 

9-23-32 


Main 


Length 
1000-1355-1680 
1002-1329-1641 
1026-1309-1602 


Table 8—Measurements of spicules of specimens of Tethya taboga (wm). Underlined numbers indicate mean values. 
1 
2 
3 


Specimens 


819 


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There are also some strongyles which mea- 
sured 490 by 16 wm, 1090 by 28.5 pm, 
1123 by 27 pm, 1268 by 25 wm and 1450 
by 30.5 wm. Several thinner styliform 
strongyloxeas are slightly sinuous. 

Megasters are spherasters, generally 40— 
60 wm in diameter, 0.3—0.6 in R/C and 16— 
20 conical rays. 

Micrasters are small tylasters in the cor- 
tex and larger oxyasters in the choanosome. 
Cortical tylasters are stout, generally 8—12 
jum in diameter with 4—8, sometimes 3, fre- 
quently irregular, microspined knobbed 
rays. Choanosomal oxyasters, generally 25— 
55 wm in diameter, with 4—8 smooth or 
spined rays, frequently bent, bifid or twist- 
ed. 

Remarks.—The identification of the 
Mexican 7. taboga specimens has been 
made by comparison with the type material. 
The two samples of Mazatlan differ slightly 
in the spheraster diameter and R/C between 
them and more considerably with the Pan- 
ama type. The megasters of the type have 
a greater size, generally 50-80 pm and a 
lower R/C (0.3-—0.5). Micrasters, tylasters 
and oxyasters, are alike in Mexican and 
type specimens. 


General remarks 


The Mexican Pacific species of Tethya 
here recorded, with the exception of TJ. fa- 
boga, represent a homogenous group, as in- 
dicated by the very similar micraster type. 
This micraster is mainly a slender stron- 
gylaster, variable, also in the same speci- 
men, from a slightly knobbed tylaster to a 
slightly tylote oxyaster. This group of spe- 
cies may be called “‘californiana”’ from the 
first described species T. californiana De 
Laubenfels, 1932 and does not show any 
evident relationship with Tethya species of 
other regions. 

Table 9 summarizes the distinctive and 
similar traits of these species as well as the 
differences with 7. taboga. This last species 
belongs to the “‘seychellensis’”’ group, char- 
acterized by two types of micrasters, corti- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


cal tylasters and larger choanosomal oxyas- 
ters. 


Acknowledgments 


This work was supported financially by 
the Instituto de Ciencias del Mar y Lim- 
nologia, Universidad Nacional Aut6énoma 
de Mexico, by CONACyT ICECXKNA- 
021996) project and by the Italian Govern- 
ment M.U.R.S.T. funds. Thanks are due to 
Dr. Martin Merino Ibarra for his support, 
Jorge Sepulveda Sanchez for the SEM mi- 
crographs, Dulce Maria Flores Sanchez for 
spicule measurement, Ignacio Palomar Mo- 
rales and Blanca Rocio Tafoya Fernandez 
for their help with image processing, Laura 
Vazquez Maldonado for the 7. ovum pho- 
tographs (Figs. 8A—B), and Jorge A. Castro 
Sanchez for the map drawing; to the anon- 
ymous reviewers whose comments im- 
proved the manuscript. 


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VOLUME 114, NUMBER 3 


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land.—Proceedings of Zoological Society of 
London: 127-141. 

Lamarck, J. B. P A. 1815. Suite des polipiers empa- 
tés.—Mémoirs de Muséum 1:69—80. 

Laubenfels, M. W. de. 1932. The marine and fresh- 
water sponges of California.—Proceedings of 
the United States National Museum 81, art. 4: 
1-140. 

. 1936. A comparison of the shallow water 

sponges near the Pacific end of the Panama Ca- 

nal with those at the Caribbean end.—Proceed- 


821 


ings of the United States National Museum 83: 
441-466. 

Molina-Cruz, A. 1986. Evoluci6n Oceanografica de la 
boca del Golfo de California.—Anales del In- 
stituto de Ciencias del Mar y Limnologia, Univ- 
ersidad Nacional Autonoma de México 13:95— 
120. 

Pallas, PR S. 1766. Elenchus zoophytorum sistens ge- 
nerum adumbrationes generaliores et specierum 
cognitarum susccinctas descriptiones cum selec- 
tis auctorum synonymis. Hagae-Comitum apud 
Petrum van Cleef, 451 pp. 

Sara, M. 1992. New-Guinean Tethya (Porifera Demos- 
pongiae) from Laing Island with description of 
three new species.—Cahiers de Biologie Marine 
33:447—467. 

. 1994. A rearrangement of the family Tethy1- 

dae (Porifera Hadromerida) with establishment 

of new genera and description of two new spe- 
cies.—Zoological Journal of the Linnean Soci- 

ety 110:355—371. 

, & G. Corriero. 1993. Redescription of Tethya 
californiana De Laubenfels as a valid species 
for Tethya aurantia var. californiana (Porifera, 
Demospongiae).—Ophelia 37:203—211. 

Thiele, J. 1905. Die Kiesel und Hornschwamme der 
Sammlung Plate.—Zoologische Jahrbticher, 
Supplement 6:107—496. 

Topsent, E. 1900. Etude monographique des spongi- 
aires de France. III. Monaxonida (Hadromeri- 
da).—Archives de Zoologie Expérimentale et 
Générale (3) 8:1-331. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(3):822. 2001. 


BIOLOGICAL SOCIETY OF WASHINGTON 
128th Annual Meeting, 29 May 2001 


President Brian FE Kensley called the 
meeting to order in the Waldo Schmitt 
Room, National Museum of Natural His- 
tory. Council members present: Roy W. 
McDiarmid (President Elect), W. Duane 
Hope, Susan L. Jewett, Rafael Lemaitre, 
(Elected Council), C. Brian Robbins (Edi- 
tor), Storrs L. Olson (Custodian of Publi- 
cations), Richard Banks (Past President), 
Janet Reid (Past President), Frank D. Fer- 
rari (Associate Editor, Invertebrates), T. 
Chad Walter (Treasurer), Carole C. Baldwin 
(Secretary). 

Minutes of the 127th Annual Meeting of 
the Society were approved, and President 
Kensley then called on Chad Walter for the 
Treasurer’s Report (Table 1). Society in- 
come for the period 1 January 2000 to 31 
December 2000 was $105,736.21; expenses 
for the same period were $123,796.96. To- 
tal assets for the Society as of 15 April 
2001 were $119,733.58, the Society’s en- 
dowment account declining by $5,013.39 
because of stock-market fluctuations. The 
Treasurer’s report was approved, and Pres- 
ident Kensley thanked Duane Hope and 
Kristian Fauchald for auditing the Treasur- 
er’s account records. 

Editor Brian Robbins then reported that 
four issues of Volume 113 of the Proceed- 
ings were published comprising 108 papers 
and 1172 pages, the largest volume in re- 


33 submissions, down from 45 in 2000. 
There continues to be no backlog for papers 
accepted in the Proceedings. 

Revision of the new guidelines for au- 
thors is nearing completion, and several op- 
tions for their publication were discussed, 
including placing them on the World Wide 
Web. The Society currently has no presence 
on the internet, but Kensley and Susan Jew- 
ett agreed to investigate development of a 
Society web site. Of considerable impor- 
tance is that the new guidelines will include 
instructions for electronic submission of 
manuscripts, which will reduce publication 
costs considerably. 

Editor Robbins announced his plans to 
move to southern California in July 2001, 
but noted that he is willing to continue 
serving as editor from his new residence. 
Details of the arrangement will be ad- 
dressed in the near future, but the Council 
agreed that Robbins’ continuing as Editor 
would provide the best chances for uninter- 
rupted production of the Proceedings. 

Following a brief discussion of who 
might replace Janet Reid as the Society’s 
representative in the Washington Academy 
of Sciences, the meeting was adjourned by 
President Kensley. 


Respectfully submitted, 
Carole C. Baldwin 


cent history. As of 1 May 2001, there were Secretary 
Table 1. Summary Financial Statement for 2000. 
General Fund Merrill Lynch Fund Total Assets 
ASSETS: JANUARY 1, 2000 42,075.82 88,746.18 130,822.00 
TOTAL RECEIPTS FOR 2000 86,496.89 19,239.32 105,736.21 
TOTAL DISBURSEMENTS FOR 2000 99,544.25 24,252.71 123,796.96 
ASSETS: DECEMBER 31, 2000 29,028.46 83,732.79 112,761.25 
NET CHANGES IN FUNDS (13,047.36) (5,013.39) (18,060.75) 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
114(3):823—824. 2001. 


INTERNATIONAL TRUST FOR ZOOLOGICAL NOMENCLATURE 


Applications published in the Bulletin of Zoological Nomenclature 


The following Applications were published on 30 March 2001 in Vol. 58, Part 1 
of the Bulletin of Zoological Nomenclature. Comment or advice on any of these 
applications is invited for publication in the Bulletin and should be sent to the 
Executive Secretary (I.C.Z.N.), c/o The Natural History Museum, Cromwell Road, 
London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk). 


Case No. 


3149 Proposed conservation of 31 species-group names originally published as 
junior primary homonyms in Buprestis Linnaeus, 1758 (Insecta, Co- 
leoptera). 

Helix lucorum Linnaeus, 1758 and Helix punctata Miller, 1774 (currently 
Otala punctata; Mollusca, Gastropoda): proposed conservation of 
usage of the specific names by the replacement of the syntypes of 
H. lucorum with a neotype. 

Ampullaria canaliculata Lamarck, 1822 (currently Pomacea canaliculata; 
Mollusca, Gastropoda): proposed conservation of the specific 
name. 

Eudorylas Aczél, 1940 (Insecta, Diptera): proposed conservation of usage by 
the designation of Pipunculus fuscipes Zetterstedt, 1844 as the type 
species. 

Halictoides dentiventris Nylander, 1848 (currently Dufourea dentiventris; 
Insecta, Hymenoptera): proposed conservation of the specific 
name. 

Parasuchus hislopi Lydekker, 1885 (Reptilia, Archosauria): proposed re- 
placement of the lectotype by a neotype. 

Euphryne obesus Baird, 1858 (Reptilia, Squamata): proposed precedence 
of the specific name over that of Sauromalus ater Duméril, 1856. 

Catalogue des mammiféres du Muséum National d’Histoire Naturelle by 
Etienne Geoffroy Saint-Hilaire (1803): proposed placement on the 
Official List of Works Approved as Available for Zoological No- 
menclature. 


824 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Opinions published in the Bulletin of Zoological Nomenclature 


The following Opinions were published on 30 March 2001 in Vol. 58, Part 1 of 
the Bulletin of Zoological Nomenclature. Copies of these Opinions can be obtained 
free of charge from the Executive Secretary, I.C.Z.N., c/o The Natural History Mu- 
seum, Cromwell Road, London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk). 


Opinion No. 


1965. Euchilus Sandberger, 1870 and Stalioa Brusina, 1870 (Mollusca, Gastro- 
poda): Bithinia deschiensiana Deshayes, 1862 and Paludina desmarestii 
Prévost, 1821 designated as the respective type species, with the conser- 
vation of Bania Brusina, 1896. 

Gnomulus Thorell, 1890 (Arachnida, Opiliones): Gnomulus sumatranus 
Thorell, 1891 designated as the type species. 

Disparalona Fryer, 1968 (Crustacea, Branchiopoda): conserved. 
Phytobius Schonherr, 1833 (nsecta, Coleoptera): placed on the Official 
List. 

Drosophila rufifrons Loew, 1873 and D. lebanonensis Wheeler, 1949 (cur- 
rently Scaptoarosophila rufifrons and S. lebanonensis; Insecta, Diptera): 
specific names conserved by the designation of a neotype for D. rufifrons. 
Odatria keithhornei Wells & Wellington, 1985 (Reptilia, Squamata): spe- 
cific name placed on the Official List. 


INFORMATION FOR CONTRIBUTORS 


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Front cover—from this issue, p. 564. 


CONTENTS 


A new species of electric ray, Narcine leoparda, from the tropical eastern Pacific Ocean (Chondrich- 

thyes: Torpediniformes: Narcinidae) Marcelo R. de Carvalho 

A new species of “Whale Catfish” (Siluriformes: Cetopsidae) from the western portions of the Amazon 

basin José Carlos de Oliveira, Richard P. Vari, and Carl J. Ferraris, Jr. 

A new species of the luvarid fish genus tAvitoluvarus (Acanthuroidei: Perciformes) from the Eocene 

of the Caucasus in southwest Russia Alexandre F. Bannikov and James C. Tyler 

Pseudothyone levini, a new species of sea cucumber (Echinodermata: Holothuroidea) from the north- 

eastern Pacific Philip Lambert and Kathryn L. Oliver 

A new leucosiid crab of the genus Nursia Leach, 1817 from Vietnam (Crustacea: Decapoda: Brachyu- 

ra) with redescription of N. mimetica Nobili, 1906 Hironori Komatsu and Masatsune Takeda 

First record of the portunid crab Arenaeus cribrarius (Lamarck, 1818) (Crustacea: Brachyura: Portuni- 

dae) in marine waters of Argentina Marcelo A. Scelzo 

First record of larvae of the rare mud shrimp Naushonia Kingsley (Crustacea: Decapoda: Laomediidae) 

from Asian waters Kooichi Konishi 

A new cavernicolous species of freshwater crab (Crustacea: Brachyura: Potamidae) from Palau Tio- 

man, peninsular Malaysia Darren C. J. Yeo 

Symbionts of the hermit crab Pagurus longicarpus Say, 1817 (Decapoda: Anomura): New observations 

from New Jersey waters and a review of all known relationships John J. McDermott 

A new species of Exosphaeroma Stebbing (Crustacea: Isopoda: Sphaeromatidae) from the Pacific coast 

of Mexico Ma. del Carmen Espinosa-Pérez and Michel E. Hendrickx 

A review of Pseudionella Shiino, 1949 (Crustacea: Isopoda: Bopyridae), with the description of a new 
species parasitic on Calcinus hermit crabs from Easter Island 

Christopher B. Boyko and Jason D. Williams 

Foliomolgus cucullus, a new genus and species of Clausidiidae (Crustacea: Copepoda: Poecilostoma- 


toida) associated with a polychaete in Korea Il-Hoi Kim 
Two new species of the Canthocamptus mirabilis group (Copepoda: Harpacticoida: Canthocamptidae) 
from South Korea Cheon Young Chang and Teruo Ishida 


Exopod and protopodial endite i of the maxillae of Sarsiellinae (Crustacea: Ostracoda: Myodocopa) 
Louis S. Kornicker 
Cypridina olimoblonga Kornicker, a new name for the junior primary homonym Cypridina oblonga 
Jones & Kirkby, 1874, and clarification of its authorship, and the authorship of the Palaeozoic gen- 
era Cypridinella, Cypridellina, Sulcuna, Rhombina, and Offa (Crustacea: Ostracoda) 
Louis S. Kornicker 
Seepiophila jonesi, a new genus and species of vestimentiferan tube worm (Annelida: Pogonophora) 
from hydrocarbon seep communities in the Gulf of Mexico 
Stephen L. Gardiner, Erin McMullin, and Charles R. Fisher 
Syllidae (Polychaeta) from San Quintin lagoon, Baja California, Mexico, with the description of a new 
genus Victoria Diaz-Castafteda and Guillermo San Martin 
Recognition of Cenogenus Chamberlin, 1919 (Polychaeta: Lumbrineridae) based on type material 
Luis F. Carrera-Parra 
On the rotifer fauna of Bermuda, including notes on the associated meiofauna and the description of a 
new species of Encentrum (Rotifera: Ploima: Dicranophoridae) Martin V. Sgrenson 
New Unguiphora (Platyhelminthes: Proseriata) from India 
Marco Curini-Galletti, Gavino Oggiano, and Marco Casu 
Studies on western Atlantic Octocorallia (Coelenterata: Anthozoa). Part 1: The genus Chrysogorgia 


Duchassaing & Michelotti, 1864 Stephen D. Cairms 
Reevaluation of Tropidopathes saliciformis Silberfeld: A hydroid originally identified as an antipathar- 
ian coral Dennis M. Opresko and Rosemarie C. Baron-Szabo 


East Pacific Mexican Tethya (Porifera: Demospongiae) with descriptions of five new species 

Michele Sara, Patricia G6mez, and Antonio Sara 
Biological Society of Washington, 128th Annual Meeting 
International Trust for Zoological Nomenclature 


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ISSN 0006-324X 


PROCEEDINGS of THE 
BIOLOGICAL SOCIETY 
or WASHINGTON 


19 DECEMBER 2001 


VOLUME 114 
NUMBER 4 


THE BIOLOGICAL SOCIETY OF WASHINGTON 
2000-2001 
Officers 


President: Brian F. Kensley _ Secretary: Carole C. Baldwin 
President-elect: Roy W. McDiarmid Treasurer: T. Chad Walter 


Elected Council 


Michael D. Carleton G. David Johnson 
W. Duane Hope Rafael Lemaitre 
Susan L. Jewett Jon Norenburg 


Custodian of Publications: Storrs L. Olson 


PROCEEDINGS 


Editor: C. Brian Robbins 


Associate Editors 


Classical Languages: Frederick M. Bayer Invertebrates: Stephen L. Gardiner 
Plants: David B. Lellinger Frank D. Ferrari 
Insects: Wayne N. Mathis Rafael Lemaitre 


Vertebrates: Gary R. Graves 
Carole C. Baldwin 


Membership in the Society is open to anyone who wishes to join. There are no prerequisites. 
Annual dues of $25.00 (for USA and non-USA addresses) include subscription to the Pro- 
ceedings of the Biological Society of Washington. Annual dues are payable on or before January 
1 of each year. Renewals received after January 1 must include a penalty charge of $3.00 for 
reinstatement. Library subscriptions to the Proceedings are: $40.00 for USA and non-USA 
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US banks, or MASTERCARD or VISA credit cards. 
Manuscripts, corrected proofs, and editorial questions should be sent to: 


EDITOR 

BIOLOGICAL SOCIETY OF WASHINGTON 
NATIONAL MUSEUM OF NATURAL HISTORY 
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Known office of publication: National Museum of Natural History, Smithsonian Institution, 
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(4):825—832. 2001. 


Merotrichous isorhiza, a nematocyst new to the Campanulariidae 


(Cnidaria: Hydrozoa), and its relevance for the 
classification of Cnidae 


Alberto Lindner and Alvaro E. Migotto 


Centro de Biologia Marinha, Universidade de Sao Paulo, Caixa Postal 83, 11600-970, 
Sao Sebastiao, SP, Brazil; and Departamento de Zoologia, Universidade de Sao Paulo, 
Sao Paulo, SP, Brazil; 

(AL) Present address: Biology Department, Duke University, Box 90338, 
Durham, North Carolina 27708, U.S.A. 


Abstract.—Merotrichous isorhizae, a nematocyst class found in only a few 
species of Cnidaria, are present in the hydroid Clytia noliformis. This nema- 
tocyst type is present in the hypostome, gonangium, and hydranth body, near 
the base of the tentacles but is absent in the tentacles themselves. The undis- 
charged capsules are similar to the B-type microbasic b-mastigophores found 
in other species of Clytia. The presence of a prominent rod in undischarged 
capsules indicates that changes may be needed in the classification of hetero- 
nemes and haplonemes, categories that embrace the great majority of nema- 


tocysts. 


Within the hydroid family Campanulari- 
idae (Hydrozoa: Leptomedusae), knowl- 
edge on the cnidome has been shown to be 
valuable for the identification of more than 
20 species (see Table | for references). In- 
formation on nematocysts has been crucial 
in solving taxonomic debates, such as the 
ones on the validity of Clytia gracilis (M. 
Sars, 1850) and Obelia longissima (Pallas, 
1766) (Ostman 1979a, 1979b, 1982; Cor- 
nelius & Ostman 1986). Two classes of 
nematocysts have thus far been reported for 
the Campanulariidae, i.e., microbasic b- 
mastigophore and holotrichous isorhiza 
(sensu Ostman 2000). These classes were 
subdivided into seven and four different 
types, respectively (Table 1), using intra- 
and interspecific size differences, as well as 
the morphology of the capsules and spines 
(Ostman 1979a, 1979b, 1982, 1988, 1999). 
However, with a few exceptions (e.g., Mig- 
otto 1996), most cnidome studies within the 
Campanulariidae were based on species 
from temperate waters (e.g., Russell 1938; 
Kubota 1976, 1978a, 1978b; Ostman 


I979a, ISTMo, IsZ, WSS, WOOD). Try 
studying the tropical species Clytia nolifor- 
mis auct., we found a nematocyst class not 
yet reported for the Campanularidae and 
present only in a few species of the Phylum 
Cnidaria: the merotrichous isorhiza. 


Materials and Methods 


Clytia noliformis was identified accord- 
ing to Calder (1991). Discharged and un- 
discharged capsules of nematocysts of C. 
noliformis were observed by squash prep- 
arations (Silveira & Migotto 1984) of living 
colonies collected along the shallow subti- 
dal coasts of Sao Sebastido (23°49.72'S, 
ASE) SS vv) mandenllinabelan(23 esse 
45°25.07'W), southeast Brazil, between 
1996-1998. Undischarged capsules of the 
proposed neotype of C. noliformis (see 
Lindner & Calder 2000), fixed in ETOH 
70%, from Castle Harbour, Bermuda, were 
also observed. Nematocysts were measured 
and photographed by light microscopy. The 
nomenclature adopted is that of Mariscal 
(1974) and Ostman (2000). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


826 


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Il ‘L ‘9 N = N odAq-g ‘adAj-v (Trl ‘ABIAT]ISORI]) Visazu1 sixXdoyji—C 
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VOLUME 114, NUMBER 4 


827 


Table 2.—Measurements of undischarged merotrichous isorhiza capsules of Clytia noliformis. 


Length (jm) 


Width (wm) Number of 


Locality {mean + SD (range) (n)] [mean + SD (range) (n)] colonies observed 
Brazil, Sao Sebastiao 21.1 = 1.6 (18.0—24.0) (37) 5.3 + 0.6 (4.0-7.0) (37) 2, 
Brazil, Whabela 20.4 + 1.1 (17.0—23.0) (26) 5.1 + 0.3 (4.5-6.0) (26) 2) 
Bermuda 19.8 = 0.6 (19.0—21.0) (10) 4.4 + 0.3 (4.0-5.0) (10) 1 


Results 


Two classes of nematocysts, i.e., micro- 
basic b-mastigophore (sensu Ostman 2000) 
and merotrichous isorhiza (sensu Mariscal 
1974), are present in the hydroid C. noli- 
formis. The microbasic b-mastigophores are 
7.1 = 0.6 wm Gmean + SD, n = 59) in 
length and 2.5 + 0.5 wm (mean + SD, n = 
59) in width. These nematocysts are abun- 
dant in the entire colony and correspond to 
the A-type described by Ostman (1979a). 
Merotrichous isorhizae are abundant in the 
hypostome, gonangium, and hydranth body, 
near the base of the tentacles but are absent 
in the tentacles. The undischarged capsule 
is elongated (ca. 20 wm in length; see Table 
2), with rounded ends. In a lateral view, one 
side is slightly convex and the other ap- 
proximately straight; a distinct axial rod, 
corresponding to the region of the armature, 
is visible inside the undischarged capsule, 
running longitudinally from its tip to the 
posterior end (Fig. 1A). The aperture is 
turned towards the straighter side. The 
everted tubule forms a 20—80° angle with 
the straighter side of the capsule (Fig. 1B). 
The region with prominent spines (arma- 
ture) is slightly shorter than one capsule- 
length (17.95 + 1.58 wm; n = 19); the dis- 
tance between its distal end and the tip of 
the capsule is about 4 capsule-lengths 
(80.68 + 8.37 wm; n = 70). The diameter 
of the tubule shortly before and after the 
armature is identical. Medusae of C. noli- 
formis have the same types of nematocysts 
so far reported for the genus Clytia (Table 
1)—microbasic b-mastigophores (A-, C- 
and D-types) and holotrichous isorhizae (1,- 
type). 


Discussion 


The A-type microbasic b-mastigophore is 
the most common nematocyst in the Cam- 
panulariidae, showing almost no variation 
in shape and size in all species studied so 
far (Ostman 1999). In contrast, the B-type 
microbasic b-mastigophore—another com- 
mon nematocyst present in all species of 
Clytia, except C. noliformis—differs in 
shape and size among species and is used 
for species identification (Ostman 1979a, 
1987, 1988, 1999; Ostman et al. 1987). Un- 
discharged capsules of the latter type are 
morphologically similar to the merotrichous 
isorhiza of C. noliformis. With light mi- 
croscopy, both nematocysts can only be dis- 
tinguished by the position of the prominent 
armature in the everted tubule. These nem- 
atocysts also have a similar distribution in 
the colony—absent from the tentacles and 
abundant in the hypostome, gonangium, 
and near the base of the tentacles (Ostman 
1979a, 1988; Lindner 2000). This suggests 
that, within the Campanulariidae, the mer- 
otrichous isorhiza may have evolved from 
microbasic b-mastigophores (or vice-versa), 
1.e., the prominent armature possibly “‘mi- 
grated”’ either from the proximal part of the 
tubule to a more distal part (microbasic > 
merotrichous) or from a distal to a proximal 
position (merotrichous — microbasic). 

Instead of merotrichous isorhizae, B-type 
microbasic b-mastigophores were reported 
for colonies identified as C. noliformis from 
Italy (Ostman et al. 1987, redefined as sB- 
type by Ostman 1987, 1999). Since these 
capsules also measured only approximately 
one-third of the length of the capsules 
found in colonies of C. noliformis from 


Fig. 1. 
A, undischarged capsule; B, discharged capsule with 
everted tubule. Scale bars: 10 wm. 


Merotrichous isorhiza of Clytia noliformis. 


Brazil and Bermuda [ca. 7 wm (see Ostman 
et al. 1987:302) and 20 wm (Table 2), re- 
spectively], it is possible that the authors 
based their observations on another species. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Although differences in cnidome may be 
indicative of different species (see Ostman 
1982, 1988), the taxonomic relevance of the 
size differences of nematocysts has not yet 
been evaluated. However, the mean size of 
the merotrichous isorhiza among different 
colonies of C. noliformis shows little vari- 
ation (Table 2). The large size may, there- 
fore, help to identify the species, and it is 
particularly important for the identification 
of fixed samples having only undischarged 
capsules. The slightly smaller size of cap- 
sules of nematocysts from Bermuda in 
comparison to those from Brazil (particu- 
larly width, ca. 16% smaller in the colony 
from Bermuda; see Table 2) is probably due 
to fixation, a procedure that causes a 15— 
20% decrease in the width of B-type mi- 
crobasic b-mastigophores in other species 
of Clytia (Lindner 2000). 

Until recently, the merotrichous isorhiza 
was a nematocyst category reported only 
for the class Hydrozoa, where it was found 
in a few species of five families of Lepto- 
medusae (Eirenidae, Eucheilotidae, Love- 
nellidae, Haleciidae and Tiaropsidae), and 
in Halitiara, a genus of Anthomedusae with 
features paralleling somewhat features 
found in the Leptomedusae (Bouillon et al. 
1988). According to Bouillon et al. (1988), 
the presence of merotrichous isorhizae sug- 
gests affinities between the families Eiren- 
idae and Eucheilotidae, as well as between 
the latter family and the Lovenellidae. 

Similarly, Boero & Sara (1987) consid- 
ered the presence of merotrichous isorhizae 
in polyps of Campalecium medusiferum 
(Torrey, 1902) and AHydranthea margarica 
(Hincks, 1862) (Haleciidae), and in medu- 
sae of Eucheilota maculata Hartlaub, 1894 
(Eucheilotidae), as a further evidence that 
the families Haleciidae and Campanulini- 
dae s.l. are phylogenetically related. Werner 
(1965), who first described this nematocyst 
class (for Eucheilota maculata), and includ- 
ed it in the category haploneme in the sys- 
tem of Weill (1934), also attributed taxo- 
nomic value to the presence of merotri- 
chous isorhizae (as ““merotriche Haplone- 


VOLUME 114, NUMBER 4 


me’’) in the cirri of medusae of Eucheilota 
and Eutima. However, the presence of mer- 
otrichous isorhizae in the scyphomedusa 
Cyanea nozakii Kishinouye, 1891 (see 
Wang & Xu 1990) and possible misidenti- 
fications of this nematocyst class as micro- 
basic or macrobasic mastigophores (as, for 
example, in Campalecium medusiferum, see 
Boero et al. 1987) indicate that merotri- 
chous isorhizae may be actually more com- 
mon among the Medusozoa. Moreover, this 
kind of nematocyst may have evolved more 
than once, and some phylogenetic inferenc- 
es based on the presence of merotrichous 
isorhizae may be misleading if not corre- 
sponding to appropriate generality levels. 
Phylogenetic relationships within the class 
Hydrozoa are poorly understood, and the 
relevance of merotrichous isorhizae for the 
systematics of the Leptomedusae (e.g., if 
they represent or not a synapomorphy of 
some taxa) can only be assessed after a 
comprehensive phylogenetic analysis. It is 
also important to emphasize that the name 
merotrichous isorhiza refers solely to fea- 
tures of the tubule of the nematocyst; other 
characters, such as a visible rod in the un- 
discharged capsule and the shape of the 
capsule, for example, should also be con- 
sidered when comparing nematocysts of 
this and other categories. 

The presence and absence of an internal 
axial rod in the undischarged capsule is a 
key character for the definition of the cat- 
egories heteronemes and haplonemes in the 
new classification of Ostman (2000). Het- 
eronemes and haplonemes were originally 
defined by Weill (1934) by the presence and 
absence, respectively, of a basally enlarged 
region of the everted tubule, the so-called 
shaft, seen in discharged capsules. In con- 
trast, Ostman (2000), in an attempt to im- 
prove the classification of Weill (1934) by 
incorporating information not available at 
his time—but without changing the nomen- 
clature—redefined heteronemes and haplo- 
nemes by the presence and absence, respec- 
tively, of a prominent rod-shaped shaft, cor- 
responding to the region of the prominent 


829 


armature, visible inside the undischarged 
capsule (regardless of the presence of an 
enlargement of the everted tubule). Since an 
internal rod is visible inside the undis- 
charged capsule of the merotrichous isor- 
hiza of C. noliformis, this nematocyst—a 
haploneme sensu Mariscal (1974)—must be 
classified as a heteroneme in the system of 
Ostman (2000). 

The presence of an internal rod (inferred 
by drawings) is also observed in the mer- 
otrichous isorhiza of, for example, Hydran- 
thea margarica (Boero & Sara 1987:133), 
Lovenella assimilis (Browne, 1905) (Hirano 
& Yamada 1985:133) and Tiaropsidium ro- 
seum (Maas, 1905) (Boero et al. 1987:296). 
However, in contrast to the classification of 
Mariscal (1974), the class merotrichous is 
not applicable in the classification of Ost- 
man (2000). Moreover, in adopting the lat- 
ter classification, the merotrichous isorhiza 
of C. noliformis may be assigned to two 
distinct classes of heteronemes, since the 
length of the proximal tubule with promi- 
nent armature is variable. Those nemato- 
cysts with the proximal tubule with prom- 
inent armature slightly shorter than 4 cap- 
sule-lengths would be classified as meso- 
basic, whereas those with the proximal 
tubule with prominent armature slightly 
longer than 4 capsule-lengths would be 
classified, according to Ostman (2000), as 
macrobasic. 

Even if the merotrichous isorhiza of C. 
noliformis could be objectively classified as 
a mesobasic or macrobasic b-mastigophore, 
we believe that this would represent a hin- 
drance, since nematocysts with distinct ar- 
matures (i.e., those with a long region with 
prominent spines starting at the base of the 
tubule and those with prominent spines 
only at a more distal position) would be 
grouped in the same categories. Since the 
merotrichous isorhiza is a nematocyst easy 
to identify and important for species iden- 
tification, such as C. noliformis, it seems 
more appropriate to maintain it as a cate- 
gory in the classification of cnidae. 

Furthermore, we believe that the visibil- 


830 


ity of an internal rod may not be a suitable 
trait to distinguish all heteronemes and hap- 
lonemes—two categories that include the 
great majority of nematocysts. This opin- 
ion, however, does not deny that further im- 
provements in the definitions of the nema- 
tocysts categories may be needed in the fu- 
ture. 


Acknowledgments 


We thank D. R. Calder and M. Zubowski 
for the loan of the proposed neotype of C. 
noliformis and L. P. de Andrade for kindly 
providing living specimens. We are also 
grateful to A. C. Marques, C. Ostman, P. 
Schuchert and W. Vervoort for their revi- 
sions and many suggestions that improved 
the manuscript. A Lindner thanks Funda¢ao 
de Amparo a Pesquisa do Estado de Sao 
Paulo (FAPESP proc. 99/00636-3) and 
Conselho Nacional de Desenvolvimento 
Cientifico e Tecnologico (CNPq) for a Mas- 
ters and an undergraduate research schol- 
arship, respectively. 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(4):833-857. 2001. 


Nipponnemertes fernaldi, a new species of swimming monostiliferous 
hoplonemertean from the San Juan Archipelago, Washington, U.S.A. 


Fumio [Iwata 


Zoological Institute, Faculty of Science, Hokkaido University, Sapporo 060-0810, Japan and 
Friday Harbor Laboratories, University of Washington, Friday Harbor, Washington 98250, U.S.A. 
(Correspondence to: Laboratory of Biology, Momijidai-Nishi 5-9-13, Atsubetsu-ku, Sapporo 004- 

0013, Japan; E-mail: fumiwata@d6.dion.ne.jp) 


Abstract.—Nipponnemertes fernaldi, new species, is described as a member 
of the order Hoplonemertea, suborder Monostilifera. In general morphology, it 
appears to be most closely related to the genus Nipponnemertes (family Cra- 
tenemertidae) and the new species is placed in this genus. The significant char- 


acters are explained and discussed. 


Among numerous nemerteans collected 
in the San Juan Archipelago, Washington in 
1964 were specimens of a monostiliferous 
hoplonemertean here described as new. 
This nemertean possesses characters that 
exclude it from any of the previously de- 
scribed species of Nipponnemertes. New in- 
formation on the morphology of monostil- 
iferans is provided with respect to: the cen- 
tral stylet-basis complex of the proboscis; 
the cerebral organs (with a sensory canal 
and a sac-like canal); a pair of middorsal 
epidermal folds in the anterior region; the 
stylet bulb of the proboscis, with a plate- 
like muscular wall and a thick layer of glan- 
dular cells; the intestinal caecum, with four 
branches or pouches and two pairs of lateral 
diverticula; the nervous system, with large 
ganglion cells in the brain; and a middorsal 
nerve extending anteriorly beyond the 
brain. 


Materials and Methods 


Two specimens were collected by dredg- 
ing, and two were collected at a night light. 
All four were fixed in Bouin’s solution (af- 
ter being anesthetized by addition of men- 
thol or 70% alcohol). Three of them were 
sectioned at 10 1m (8 pm for the proboscis 
apparatus of the holotype), and stained with 


Delafield’s hematoxylin and eosin. An 
Olympus or Sony video camera with Sym- 
phonic monitor television and Sony color 
video printer, in addition to an Olympus 
BH-2 microscope, were used for observa- 
tion of the material. 


Nipponnemertes fernaldi, new species 
Figs. 1-9, Tables 1, 2 


Generic diagnosis.—Monostiliferous ho- 
plonemerteans with rhynchocoel nearly 
equal to body length, proboscis sheath com- 
posed of a wickerwork of interwoven cir- 
cular and longitudinal muscles; body wall 
musculature, especially longitudinal layer, 
well developed, with layer of diagonal mus- 
cles between the circular and the longitu- 
dinal muscles, longitudinal musculature not 
anteriorly divided; stomach with a small di- 
verticulum or caecal folds; intestinal cae- 
cum with short anterior diverticula and with 
lateral diverticula; cerebral sensory organ 
large and extending behind the dorsal ce- 
rebral ganglia; nervous system without neu- 
rochords and neurochord cells; accessory 
lateral nerves absent; blood vascular system 
with three longitudinal vessels, mid-dorsal 
vessel with single vascular plug; frontal or- 
gan present; cephalic glands well developed 
but not reaching posteriorly behind the ce- 


834 


rebral ganglia, except as lateral bands; ex- 
cretory system extending from rear of brain 
to hind regions of foregut; sexes separate. 
Specific diagnosis.—Pale brown, without 
pattern; with two pairs of oblique cephalic 
grooves and dorsomedial longitudinal 
grooves; each anterior oblique cephalic 
groove is subdivided by 21 transverse sec- 
ondary grooves, 14 dorsal to the opening of 
cerebral organ canal and 7 on the ventro- 
lateral surface of the head below the canal 
opening; paired posterior oblique cephalic 
grooves originating at the end of the dor- 
somedial longitudinal grooves; dorsomedial 
longitudinal grooves forming anteriorly a 
pair of epidermal folds; with numerous 
large ocelli (76 in the holotype, 78 in par- 
atype 1, and 52 in paratype 3) in four 
groups; dermis thick; rhynchodeum provid- 
ed with a circular muscle layer; precerebral 
septum is of the dissolved type composed 
of separate muscular bundles; proboscis di- 
aphragm provided with central stylet-basis 
complex, which includes a basis consisting 
of ovoid body to which the central stylet is 
attached, columnar structure (with oblique- 
ly oriented projection) posterodorsal to it, 
and more massive posterior portion resting 
on muscular bolster; stylet bulb with a 
plate-like muscular wall and a thick layer 
of glandular cells; 2 or 3 accessory stylet 
pouches; proboscis with 14 nerves; mouth 
small and located on the ventral side of the 
rhynchodeum; esophagus, anterior to brain, 
surrounded by longitudinal muscle; intesti- 
nal caecum with anterior branches or 
pouches and two pairs of lateral diverticula; 
intestinal diverticula branched; brain with 
neurilemma and large ganglion cells; mid- 
dorsal nerve extending anteriorly beyond 
the brain; lateral nerve cords with myofi- 
brillae and large efferent nerves; ventral 
ganglia not distinctly separated from the 
dorsal ganglia except posteriorly; dorsal 
ganglia without bifurcated fiber core; paired 
midgut, foregut, and esophagus nerves orig- 
inating from the ventral brain; cerebral sen- 
sory organ with neurilemma, the cerebral 
organ canal, located at the anterior end of 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the cerebral sensory organ, branching into 
the medial sensory canal and lateral sac-like 
canal with two kinds of glandular masses; 
glands lacking at the bifurcation of the ce- 
rebral organ canal; excretory pores lateral 
to the nerve cords and situated in the pos- 
terior portion of the nephridial region. 

Type material.—Holotype (USNM 
1000131), paratype 1 (USNM_ 1000132), 
paratype 2 (USNM 1000133) and paratype 
3 (USNM 1000134) are deposited at the 
National Museum of Natural History, 
Smithsonian Institution, Washington, DC. 

The holotype (female) consists mostly of 
a set of serial sections mounted on 81 
slides. These include transverse sections of 
the anterior portion of the body, midgut, 
tail, and proboscis, and frontal sections of 
the proboscis apparatus. The remainder of 
the holotype consists of short pieces that 
were not sectioned. Paratype 1 (male) con- 
sists of serial sections mounted on 76 
slides. These include transverse sections of 
the anterior portion of the body, midgut, 
tail, and proboscis, and frontal sections of 
the proboscis apparatus. Four pieces were 
not sectioned. Paratype 2 (male) consists of 
serial sections mounted on 29 slides, which 
include horizontal sections of the anterior 
portion of the body and proboscis appara- 
tus, and transverse sections of the tail and 
two pieces that were not sectioned. Para- 
type 3 was not sectioned (it dried once after 
it was preserved). 

Type locality.—Holotype: Peavine Pass, 
between Blakely Island and Obstruction Is- 
land, Washington, U.S.A. (48°34.7'N, 
122°50.5'W); collected on 21 Sep 1964 by 
dredging, gravel bottom, depth between 42 
and 56 m. Paratypes | and 2: Friday Harbor, 
Washington, U.S.A.; collected by night 
light in water column, depth 4 m on 12 
Sep 1964. Paratype 3: Willow Island 
(48°32.4'N), Washington, U.S.A.; collected 
on 10 Sep 1964, gravel bottom, depth be- 
tween 18 and 33 m. 

Etymology.—The species name com- 
memorates the late Professor Robert Fer- 
nald, former Director of Friday Harbor Lab- 


VOLUME 114, NUMBER 4 


oratories of the University of Washington, 
who collected two specimens (Paratypes | 
and 2). 


Description 


External features.—The holotype is 5 cm 
long and 6 mm wide, but 8 mm wide while 
moving (Fig. la, b); paratype 1 is 8 cm long 
and 7 mm wide anteriorly and 10 mm wide 
in the midgut region (Fig. Ic, d); paratype 
2 is 6 cm long and 8 mm wide; paratype 3 
is 4 cm long and 10 mm wide, but 7 cm 
long and 5 mm wide when narcotized (Fig. 
1h—j). The extruded proboscis of the holo- 
type is 2.7 cm long (Fig. la). The probos- 
cises of paratypes 1—3 were not extruded 
when the specimens were preserved. The 
proboscises of paratypes 1 and 2 were re- 
moved from the body for observation of the 
proboscis apparatus (Fig. 1d). The head is 
rounded on the anterior margin and slightly 
separated from the body by a constriction 
formed by the anterior oblique cephalic 
grooves. 

In the living state, the body is flattened 
dorsoventrally, convex on the dorsal side, 
tapering posteriorly to a blunt tip. The color 
is pale brown on the dorsal surface, and 
more pale brown on the ventral side, with 
a colorless lateral margin. In the middorsal 
portion of the body, the proboscis extends 
anteroposteriorly within the large rhyncho- 
coel (Fig. 1b). The brain is recognizable 
from above as a pair of reddish markings 
Gage lesh): 

On the dorsal side of the head there are 
two pairs of oblique cephalic grooves (Fig. 
le, f, h, i). The anterior grooves extend 
around the margins of the head end and are 
continued on the ventral surface; they each 
have 21 short secondary grooves perpen- 
dicular to them; 14 of these secondary 
grooves lie dorsal to the opening of the ce- 
rebral organ canal (Fig. 11) and 7 short sec- 
ondary grooves lie below the cerebral organ 
canal opening on the ventrolateral surface 
of the head. The posterior oblique cephalic 
grooves are limited to the dorsal side of the 


835 


body, and together they form a V-shaped 
pattern (Fig. le, f, h). Two closely parallel 
longitudinal grooves are located on the 
mid-dorsal part of the head, as in some spe- 
cies of Nipponnemertes and other cratene- 
mertids. They intersect the anterior oblique 
cephalic grooves and extend to the posterior 
oblique cephalic grooves (Fig. le, f, h, 1). 
Their course is actually longer than shown 
in the figures, but their most anterior por- 
tions, which extend to the tip of the head, 
are evident only in transverse sections. In 
dorsal view, the anterior portion of each 
dorsomedial longitudinal cephalic groove 
has a median darkened line due to an epi- 
dermal depression that exhibits double epi- 
dermal folds in transverse sections (Figs. 
le, ie Ja, 22). 

The rhynchodeal opening is a slender 
lengthwise pore on the anteroventral por- 
tion of the head (Fig. 1g, j). In both the 
holotype and paratype 1, the right and left 
sides of the head region each have up to 39 
large ocelli. 

Body wall, musculature, and parenchy- 
ma.—The epidermis is of uniform thickness 
both in the cephalic region (average 110 
um) and in the midgut region (average 50 
ym). Large unicellular glands are distrib- 
uted over the entire body. Small, slender 
cyanophilous glands are embedded between 
the large glands. 

Two middorsal epidermal folds in the an- 
terior region are separated by a deep me- 
dian groove (Fig. 2a), but posteriorly the 
folds unite and the groove disappears. The 
epidermis of the folds has only a few cya- 
nophilous glands. The secondary grooves 
of the anterior oblique cephalic grooves, 
which run perpendicular to the anterior 
grooves, cut deeply into the epidermis and 
are distinguished by a thin epithelium free 
of glands, and also by comparatively long 
cilia (Fig. 2b). 

The dermis, 20 pm to 150 pm thick, is 
from one-third to three times the thickness 
of the epidermis. It has fibrils that are pre- 
dominantly circular, and contains numerous 
cells with nuclei approximately 4 wm in di- 


836 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


h I 

Fig. 1. Nipponnemertes fernaldi, new species, external features. (a) holotype, dorsal view; (b) sketch of 
holotype, dorsal view; (c) paratype 1; (d) paratype 1, and proboscis apparatus; (e) sketch of head of holotype, 
dorsal view; (f, g) sketches of head of paratype 1 in dorsal and ventral views; (h—j), sketches of head of paratype 
3 in dorsal, lateral, and ventral views. Scale = 1 cm (a, d). ac, anterior chamber of proboscis; ah, anterior 
oblique cephalic groove; br, brain; he, hind end of proboscis; lg, longitudinal cephalic groove; oc, opening of 
cerebral organ; ol, ocelli behind anterior oblique cephalic groove; ph, posterior oblique cephalic groove; pp, 
posterior portion of proboscis; ro, rhynchodeal opening; sg, secondary groove; sr, stylet region of proboscis. 


VOLUME 114, NUMBER 4 


ameter (Fig. 2c). Small nerves are found in 
the dermis. 

The body-wall musculature consists of an 
outer circular layer, inner longitudinal layer, 
and a recognizable diagonal muscle layer 
between them (Fig. 2d) (cf. Crandall 
1993a). The circular and longitudinal mus- 
cle layers extend to the tip of the head, but 
decrease in thickness precerebrally. The cir- 
cular layer posterior to the brain region has 
a fairly uniform thickness of 60 wm. The 
longitudinal muscle layer consists of bun- 
dles arranged perpendicular to the body sur- 
face; the dorsal and ventral bundles become 
thicker (up to 550 ym thick) in the stomach 
region (Fig. 2e). Longitudinal muscle fibers 
embedded in the parenchyma of the precer- 
ebral region originate from the inner por- 
tion of the longitudinal body-wall muscu- 
lature and extend to the tip of the head. In 
the precerebral and cerebral region, dorso- 
ventral and radial muscle fibers are not pre- 
sent. Behind the cerebral sensory organs, 
dorsoventral muscle fibers appear in the pa- 
renchyma between the proboscis sheath and 
the lateral longitudinal body-wall muscula- 
ture. In the midgut region, well-developed 
dorsoventral musculature, consisting of 
crowded fibers, is found between the intes- 
tinal diverticula (Fig. 2f). Parenchyma is 
clearly evident along the lateral sides of the 
body in the intestinal region. 

Rhynchodeum, rhynchocoel, and probos- 
cis.—The precerebral region, anterior to the 
opening of the rhynchodeum, forms a short 
snout, which is flattened dorsoventrally and 
contains the ocelli and frontal glands asso- 
ciated with the frontal organ (Fig. 2g). A 
midventral epidermal groove leads from the 
rhynchodeal opening to the tip of the snout 
(Fig. 2g, h). The lining of the groove, form- 
ing a narrow median depression, does not 
differ from the epidermis of the head re- 
gion, except for a boundary area around the 
opening of the rhynchodeum, where an ep- 
ithelium with cilia but without glands is 
present. The dermis and musculature dorsal 
to the groove are continuous with those of 
the remainder of the body wall but are thin- 


837 


ner. The longitudinal musculature, however, 
is displaced by the cephalic glands (Fig. 3a, 
b). 

In paratype 1, the precerebral region does 
not form a short flattened snout and the 
mid-ventral groove runs from the tip of the 
head to the rhynchodeal opening. 

The rhynchodeum is lined entirely with 
a ciliated epithelium without a glandular 
content. The epithelium rests on a layer of 
circular musculature 10—S5O ym thick in the 
holotype, 15-150 wm thick in paratype 1; 
the thickness of the the layer of circular 
musculature gradually increases toward the 
posterior end of the rhynchodeum. In the 
holotype, whose proboscis was extruded, 
the rhynchodeum is cylindrical and has a 
length of about 500 p.m. Its diameter at the 
opening measures 350 pm; at the posterior 
end it measures 450 pm. At the posterior 
end of the rhynchodeum, the circular mus- 
cle layer around this cavity, properly called 
the rhynchodeal sphincter, is 50 wm thick; 
the inner circular muscle layer of the pro- 
boscis fuses with it, resulting in compres- 
sion of the space of the rhynchodeum (Fig. 
3b). Part of the fused circular muscle layer 
measures 120 wm thick, but the thickness 
decreases posteriorly for about 200 wm, 
where the fused layer extends transversely 
to join the circular muscle layer of the body 
wall (Fig. 3c, d). The inner longitudinal 
muscle layer of the proboscis, within which 
the proboscis nerves are situated, becomes 
thicker posteriorly and forms the precere- 
bral septum of the proboscis apparatus (Fig. 
3d, e). Thus the proboscis insertion and the 
precerebral septum are distinct. 

In paratype 1, in which the proboscis was 
not extruded, the precerebral septum of the 
proboscis apparatus consists of about 20 
loosely arranged bundles of longitudinal 
muscle. Farther posteriorly, the longitudinal 
muscles closely surround the thick circular 
muscle layer around the rhynchodeum. 

Posterior to the precerebral septum, a 
layer about 20 pm thick (Figs. 3e, f, 4c, 7e), 
in which circular and longitudinal muscu- 


838 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Nipponnemerytes fernaldi, new species. Transverse sections of anterior and middle portions of 
body, showing (a) two epidermal folds (separated by a central depression) between longitudinal cephalic 
grooves and a central depression, (b) secondary grooves of anterior oblique cephalic groove (right side), (c) 
dermis, (d) middorsal nerve and diagonal muscle layer, (e) stomach, (f) midgut and lateral diverticula of 
midgut, (g) epidermal ciliated groove leading to rhynchodeum, and (h) pit of frontal organ. Scale = 0.5 mm 
(e, f), 200 pm (b, g), 100 pm (a), 50 pm (c, d). Sections 5—204 from anterior end of body (a-e, g, h); section 
288 from anterior end of middle portion of body (f). cd, cell in dermis; ce, cephalic glands; cg, ciliated 
groove; cm, circular muscle layer of body wall; cw, central depression between mediodorsal longitudinal 
cephalic grooves; dm, diagonal muscle layer; do, dorsoventral muscle; dv, dorsal blood vessel; fg, frontal 
gland; fo, frontal organ; Id, lateral diverticulum of midgut; lg, longitudinal cephalic grooves; Im, longitudinal 


VOLUME 114, NUMBER 4 


lature are interwoven, underlies the epithe- 
lium of the proboscis sheath. 

In paratype 1, the rhynchodeum and 
esophagus, situated below the cephalic 
glands, are surrounded by thick bundles of 
longitudinal musculature and a thin layer of 
circular muscles. 

In the area in which the circular muscle 
layer of the rhynchodeum and that of the 
proboscis are united into a thick layer, a co- 
agulated mass of protein-rich rhynchocoel 
fluid, about 120 pm thick, appears within 
the proboscis (Fig. 3b, c). The posterior part 
of this coagulated mass is hollowed out; the 
space marks the beginning of the rhyncho- 
coel (Fig. 3d, e). The wall of the mass 
around the space becomes narrower and 
comes close to the epithelium of the pro- 
boscis sheath (Fig. 3g). In paratypes 1 and 
2, a mass of coagulated rhynchocoel fluid 
was not observed (the proboscis was inside 
the body at the time of preservation). 

The rhynchocoel within the proboscis 
sheath ends a little anterior to the posterior 
nerve anastomosis. It is narrow in the fore- 
gut region, where the longitudinal muscu- 
lature of the body wall is well developed. 
The pressure of peripheral tissues causes 
the rhynchocoel walls to buckle inward into 
various folds that protrude into the lumen 
of the rhynchocoel, forming a pair of dorsal 
and ventral ridges (Figs. 3g, h, 4a, b). In 
the intestinal region, the rhynchocoel wid- 
ens and becomes cylindrical (Fig. 4c). 

In paratype 1, a pair of small dorsal ridg- 
es of the rhynchocoel are present in the ce- 
rebral and stomach regions, and a pair of 
deep ventral ridges also are present at the 
lateroventral corners of the rhynchocoel. 

The structures associated with the central 
stylet form an unusual complex, as yet not 
reported for any monostiliferan hoplone- 
mertean. The following observations are 


< 


839 


based on sections of the holotype (Fig. 5a— 
o). The stylet itself, about 10 pm long, is 
attached to an ovoid structure in which 
many small cellular or granular bodies are 
embedded (Figs. 4d, 5c—e, 6a). This struc- 
ture, about 25 wm long and 17 pm wide, 
has a thick, slightly irregular wall. Postero- 
dorsal to it is a nearly columnar, eosinoph- 
ilous stalk bearing a projection that is ori- 
ented obliquely; this has a darkened edge 
on one side (Figs. 4f, 5g, h, 6a). The stalk 
is essentially a prolongation of the more 
massive posterior portion of the basis, about 
110 pm long by 80 pm wide. Most of this 
is stained pale purple, but the nearly conical 
anterior half of it has a thick coating that is 
deeply stained purple and has sharp or blunt 
protuberances, as well as a collar-like ridge 
that encircles the base of the columnar stalk 
(Figs. 4f, 5h). The conical portion is free 
from the tissue surrounding it, but the pos- 
terior bulbous part is tightly bound to tissue 
(Fig. 5f—n). 

In paratype 1, the tip of the basis is 
curved downward and the anterior portion 
of the central stylet, 17 wm long, is dam- 
aged (Fig. 4g). Several large cellular bodies 
and a few deformed cyanophilous structures 
are present between the central stylet and 
the basis. The basis is conical, and its lateral 
surfaces, stained purplish, have several 
small and irregular protuberances. One side 
of the basis faces the ductus ejaculatorius 
(Stricker & Cloney 1981). In paratype 2, 
however, the tip of the basis is not curved, 
and a rectangular body containing a mass 
of small cyanophilous spheres rests on the 
tip of the conical portion of the basis (Fig. 
4h). The rectangular body, with a wall 5 wm 
thick, measures 32 jm wide and 37 pm 
high. The basis is conical, as in the holo- 
type; the free portion of its dorsal side has 


muscle layer of body wall; ls, most lateral groove on lateral side of head; md, middorsal nerve; mi, midgut; ps, 
posterior end of cerebral sensory organ; sg, secondary groove; sm, submuscular gland; st, stomach; vp, vascular 


plug inside proboscis sheath. 


840 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Nipponnemertes fernaldi, new species. Transverse sections of anterior portion of body, showing (a) 
flattened portion of esophagus under rhynchodeum, (b) tight connection of circular muscle layer of rhynchodeum 
to that of proboscis, (c) united circular muscle layers of rhynchodeum and proboscis, (d) reduction of thickness 
of united circular muscle layers running laterally to circular muscle layer of body wall, (e) formation of pre- 
cerebral septum from longitudinal muscle layer of proboscis, (f) enlarged proboscis sheath formed from epithe- 
lium of proboscis, and (g, h) successive changes in shape of rhynchocoel. Scale = 0.5 mm (g, h), 200 ~m (a— 
e) and 50 pm (f). 46-372, numbers of sections from anterior end of body. ar, artifact; ce, cephalic glands; ci, 
circular muscle layer of rhynchodeum; cl, cephalic blood lacuna; cp, circular muscle layer of proboscis; cr, 
circular muscle layer of rhynchodeum united to that of proboscis; dr, dorsal ridge of proboscis sheath; dv, dorsal 
blood vessel; es, esophagus; lp, longitudinal muscle layer of proboscis; ms, membranous sheath of coagulated 


VOLUME 114, NUMBER 4 


purplish edges, but lacks protuberances. In 
paratype 2, the central stylet is lacking. 

The basis rests on a large bolster con- 
sisting of interwoven circular and longitu- 
dinal muscle (Fig. 4e). It is 110 wm long 
medially, 150 wm long laterally, and 370 
um wide. The basis is surrounded by cir- 
cular and longitudinal muscle bundles ex- 
tending dorsally from the bolster. This mus- 
cular arrangement continues beyond the ba- 
sis to the middle area of the stylet column, 
which extends from the tip of the proboscis 
to the central stylet of the proboscis appa- 
ratus (Fig. 4e). Posterior to the basis, the 
bolster is divided into two parts, with a 
space between. The bolster, basis, and mus- 
cular wall are continuous with the wall of 
the middle part of the proboscis. The clus- 
ters of eosinophilous glands of the probos- 
cis epithelium are evident at the anterior 
end of this organ (Fig. 4e). 

Posterior to the bolster, there is a large 
bulb-like structure called the dilated am- 
pulla or stylet bulb (Stricker 1985) (Fig. 
4e); its wall is about 50 wm thick, and con- 
sists of interlacing circular muscles, thick 
connective tissue, a plate-like muscular wall 
on its anterior side, and a thick layer of 
glandular cells. In Nipponnemertes punc- 
tatulus and N. bimaculatus (reported by 
Iwata 1951, 1954), the platelike muscular 
wall and the thick layer of glandular cells 
were not found. In the area where a space 
is formed between the two halves of the 
divided bolster, the muscular anterior plate 
of the bulb is extended forward, and a nar- 
row duct called the ductus ejaculatorius 
connects the interior of the stylet bulb with 
the space. On the posterior side of the bulb, 
a narrow canal leads to the posterior cham- 
ber of the proboscis (Fig. 7a). 

In the holotype and paratype 1, two large 
pouches of accessory stylets are situated an- 


<< 


841 


terior to the central stylet, between the outer 
epithelium and the muscular wall of the 
bolster (Figs. 4e, 5a). Through a narrow 
curved duct, each pouch, measuring 265 
wm dorsoventrally, 230 wm long, and 170 
lum wide, opens into the lumen of a canal 
that is directed upward from the basis. In 
each pouch, there are four accessory stylets, 
about 10 um long; they lie in vesicles about 
30 pm by 20 pm (Figs. 4e, 5a). In paratype 
2, there are three pouches of accessory sty- 
lets, and the number of stylets is 6, 8, and 
8, respectively. 

The proboscis has an outer circular and 
inner longitudinal muscle layer (Fig. 3a, c); 
there are 14 nerves within the proboscis in 
the holotype and paratype 1. 

Alimentary canal.—The alimentary canal 
has five major divisions: esophagus, stom- 
ach, pylorus, midgut (with anteriorly di- 
rected caecum and lateral diverticula), and 
hindgut. The mouth is small and on the 
ventral side of the head close to the rhyn- 
chodeal opening (Fig. 7b). The distance be- 
tween the rhynchodeal opening and the 
mouth is 150 pm in the holotype, 200 wm 
in paratype 1. Neither the esophagus nor the 
stomach has longitudinal or circular mus- 
culature; the shape of the esophagus is in- 
fluenced by the longitudinal musculature of 
the body wall, situated on its dorsal and lat- 
eral sides. The esophagus soon flattens dor- 
soventrally (Fig. 3a). At first its epithelium 
is very thin but then thickens. It is not cil- 
iated, but middorsally it has a mass of gland 
cells projecting inward. Near the level of 
the midventral portion of the brain, the 
esophagus widens and develops two folds; 
these mark the transition into the stomach. 
The first fold incorporates some of the glan- 
dular cells. The stomach epithelium, in 
which eosinophilous gland cells begin to 
predominate, has two folds that are contin- 


fluid; pc, precerebral septum of proboscis apparatus; pn, proboscis nerves; pr, proboscis in rhynchodeum; pw, 
proboscis sheath, consisting of interwoven circular and longitudinal muscles; rc, rhynchocoel; rd, rhnynchodeum; 


vr, ventral ridge of proboscis sheath. 


842 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Nipponnemertes fernaldi, new species. Transverse sections of anterior and middle portions of body, 
showing (a—c) successive changes in shape of rhynchocoel, (d) central stylet on ovoid anterior portion of basis 
in the holotype specimen, (e) columnar portion of basis in the holotype specimen, (f) central stylet damaged in 
shape in the specimen of paratype 1, (g) rectangular anterior portion of basis in the specimen of paratype 2, (h) 
anterior end of proboscis extruded outside, showing proboscis apparatus in the holotype specimen. Scale = 0.5 
mm (a-—c, e), 200 wm (f, g) and 50 wm (d, fh). 3, 8, 38, and 67, number of sections of proboscis apparatus (d— 
h); 512 and 607, numbers of sections from anterior end of body (a, b); 211, section from middle portion of 
body (c). ao, anterior end of ovaries; bl, bulb-like basis of central stylet; bs, bolster; ca, circular muscle bundle 
of proboscis sheath; cc, cover of stylet; ci, collar-like ridge of massive posterior portion of basis (the ridge 
encircles the columnar portion of the basis); co, nearly conical part of posterior portion of basis, free of sur- 


VOLUME 114, NUMBER 4 


uous with those of the esophagus, as in 
most species of Nipponnemertes. At the 
level of the posterior end of the cerebral 
sensory organs, the stomach enlarges and 
develops a third fold (Fig. 2e). Over most 
of its length, the stomach is more than twice 
as wide as it is where it joins the esophagus 
(Fig. 2e). Beneath the stomach is a substan- 
tial mass of longitudinal body-wall muscle 
(Fig. 2e). Measured from the anterodorsal 
to posteroveniral limits of the gastric folds, 
the stomach is about three times as long as 
the brain. 

In the holotype, the pylorus, which fol- 
lows the stomach, is about 1.5 mm long and 
about two and one-half times as long as the 
stomach. It originates from the right side of 
the stomach, between its lateral wall and its 
principal fold. The left side of the stomach 
continues posteriorly into a diverticulum 
formed as an artifact of contraction (Fig. 
7c). The pylorus is oval in transverse sec- 
tion; it is about 0.25 mm wide at its junc- 
tion with the stomach, and narrows to 0.15 
mm at the point where it opens into the 
midgut (Fig. 4b). Its ciliated epithelium 
contains cyanophilous gland cells and is 
thus different from the lining of the poster- 
odorsal area of the gastric cavity (Fig. 7d). 

In paratype 1, the pylorus is about 4 mm 
long and about three times as long as the 
stomach. The diverticulum of the stomach, 
about 0.6 mm long, has three caecal folds 
in its outer portion. 

In the holotype, the intestinal caecum has 
four anterior tubular branches, of which 
three arise from the main caecum (Fig. 6b). 
Posteriorly, it has two pairs of lateral di- 
verticula; those of the anterior pair are of 
unequal length and are parallel to the main 
caecum. Those of the posterior pair form a 
tube connected by a short duct to the main 


<_ 


843 


caecum; this tube extends posteriorly be- 
yond the pylorus. The intestinal caecum and 
anterior branches extend anteriorly for 
about half the length of the pylorus. 

In paratype 1, the intestinal caecum has 
four anterior pouches that originate from 
the main caecum just anterior to where the 
first pair of lateral diverticula branch off; 
the main caecum does not extend any far- 
ther anteriorly. The intestinal caecum, about 
1.5 mm long, extends anteriorly above the 
posterior portion of the pylorus for less than 
half its length. 

Most of the lateral diverticula of the an- 
terior part of the intestine are divided into 
dorsal and ventral lobes (Fig. 2f); the lobes 
are unequal and variable in number and 
length. The epithelium of the intestine con- 
tains eosinophilous gland cells. In the ho- 
lotype, this part of the digestive tract, cir- 
cular in transverse sections, does not have 
its own musculature, but in paratype 1, 
there are a few circular muscles in the wall 
of the intestine. 

In paratype 2, the anus opens ventrally 
near the posterior end of the body. It is a 
slit about 100 pm long, and has an inside 
diameter of 30 wm transversely and 15 wm 
dorsoventrally. The wall is 40 wm thick. 
The rectum is 140 pm long and is surround- 
ed by a circular muscle layer 50 wm thick. 
In paratype 1, the rectum has the same 
structure as in paratype 2. In the holotype, 
the series of sections of the posterior por- 
tion of the body is not complete. 

Blood-vascular system.—The blood vas- 
cular system has three longitudinal vessels. 
The two cephalic blood vessels lateral to 
the rhynchodeum anastomose above the 
rhynchodeum near the tip of the head to 
form the cephalic loop. Farther posteriorly, 
however, the cephalic vessels become large 


rounding tissue; cp, columnar portion of basis; cs, central stylet; ct, bulbous part of posterior portion of basis, 
tightly bound to tissue; cu, intestinal caecum; dv, dorsal blood vessel; la, longitudinal muscle bundle of proboscis 
sheath; Id, lateral diverticula of midgut; op, opening of pylorus; or, ovary; ov, ovoid anterior portion of basis; 
pa, pouch of accessory stylet; pm, platelike muscular wall; py, pylorus; tc, dense connective tissue; tg, dense 
layer of glandular cells; wi, stylet bulb-wall consisting of circular muscle. 


844 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 5. Nipponnemertes fernaldi, new species. Successive transverse sections of proboscis apparatus in the 
holotype specimen (a—o), showing especially accessory stylet (a), bulbous proboscis vesicle (c—e), and obliquely 
oriented projection of columnar portion of basis (f-1). Scale = 150 ym, 1-16, numbers of sections of proboscis 
apparatus. as, accessory stylet; ct, bulbous part of posterior portion of basis; pa, pouch of accessory stylets. 


VOLUME 114, NUMBER 4 


lacunae that turn medially in front of the 
brain. At the level of the anterior end of the 
ventral ganglia, they become narrowed into 
vessels that enter the brain ring without giv- 
ing off cerebral vessels and then run along- 
side the rhynchocoel (Fig. 3g). In the ho- 
lotype, the right vessel soon communicates 
with the dorsal vessel, which then enters the 
proboscis sheath, at the level of the poste- 
rior end of the brain, as a single median 
vascular plug, and extends medioventrally 
in the sheath as far as the anterior pyloric 
region, a distance of about 1.4 mm (Figs. 
2e, 7c—e, g, 8e). This portion of the vascular 
plug seems to form a specialized commu- 
nication with the rhynchocoel (Fig. 7e). Af- 
ter passing ventrally out of the rhynchocoel, 
the dorsal vessel continues posteriorly un- 
der the proboscis sheath between the rhyn- 
chocoel and the alimentary canal (Figs. 2f, 
3h, 4a, b). In paratype 1, however, it is the 
left vessel that communicates with the dor- 
sal vessel at the level of the posterior por- 
tion of the cerebral sensory organs or be- 
hind the brain, and the single median vas- 
cular plug extends as far as the middle py- 
loric region, a distance of about 3.1 mm. 

Behind the cerebral organs, the lateral 
vessels enter the nephridial region, which 
extends from the posterior end of the ce- 
rebral organs to the level of the anterior 
portion of the pylorus (in the holotype) or 
to the level of the anterior portion of the 
intestine (in paratype 1) (Figs. 4a, 7c, d, f); 
there the vessels are dorsal to the lateral 
nerve cords. 

Transverse anastomoses of the lateral 
vessels in the intestinal region were not ob- 
served. 

In both paratype 1 and 2, the dorsal and 
lateral vessels anastomose just above the 
posterior end of the rectum, where a com- 
missure of the lateral nerves and the end of 
the rhynchocoel come together at the same 
transverse level. 

Nervous system.—The brain is situated 
immediately behind the large cephalic la- 
cunae. The right lacuna is 340 pm wide and 
100 wm high. The dorsal and ventral gan- 


845 


glia are not demarcated externally. The ven- 
tral commissure of the brain is situated near 
the anterior end of the brain and the ventral 
ganglia are not distinctly separated from the 
dorsal ganglia, except posteriorly. They be- 
come completely separated for a distance of 
30 pm. (Figs. 6c, 7g). The dorsal and ven- 
tral fiber cores are not divided anterior to 
the middle portion of the brain; the dorsal 
cores are more voluminous than the ventral 
cores. The dorsal ganglia do not extend as 
far posteriorly as the ventral ganglia, and 
dorsal and ventral lobes are lacking (Fig. 
6c). A thick nerve cord from the posterior 
end of each dorsal ganglion enters the cor- 
responding cerebral sensory organ (Figs. 
6c, 7g). A long, thin dorsal commissure (40 
wzm thick), and a short, much thicker ventral 
commissure (110 pm thick), connect the 
right and left ganglia at about the same 
transverse level (Fig. 6c). In paratype 1, 
however, the dorsal commissure (70 wm 
thick) is situated a short distance (30 pm) 
farther anterior than the ventral one (130 
yum). The dorsal commissure curves up- 
ward to lie above the dorsal ridge of the 
proboscis sheath. On both sides, the brain 
is in contact with the dorsolateral wall of 
the esophagus, which is spanned dorsally 
by the ventral commissure. The brain and 
lateral nerves are covered by a thin neuri- 
lemma and a thick layer of fibrous connec- 
tive tissue (Fig. 7h). In the intestinal region, 
however, the neurilemma of the lateral 
nerves is not evident. In the cerebral fiber 
core of each dorsal ganglion, there are nu- 
merous large cellular bodies (Fig. 8a, b), 
similar to those in Nipponnemertes punc- 
tatulus (reported by Iwata 1951) (Fig. 9c, 
d) and the reptantic polystiliferan Kamegi- 
nemertes parmiornatus (Iwata 1998). The 
structure of these cellular bodies is not 
clearly defined. 

In the mass formed by the dorsal and 
ventral ganglia, there is, in the outer part of 
the right side of the brain, a large ganglion 
cell 20 wm long; its nucleus measures 6 
wm. It is situated immediately behind the 
ventral commissure (Fig. 8c). A large gan- 


846 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


cs cc 


OV 
wt Oy . td wea 
ety e ort oy. . , 
ap Ri panc canes oo cl 3 
ey umemaliesip ts 
welts eee B | é bm 
centers ee co B Z 
Cee ee a: 
aoa ae ; 4 
oF > A SY ee j 
Di Ne ee py y BES 
: : a ce ~ ne i 
< } 
» ¢ 4 
?, 
| 
j 
FY fl 
t 
7 
: i 
a SOG AG oe aS Wi 
“No FS We ieme ven en K 
oy “he 8S ree ERE , 
a “pes © wn oe 
} cu 
2 
) 
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C 


Fig. 6. Nipponnemertes fernaldi, new species. (a) Drawing of frontal sections of central stylet-basis complex 
of proboscis, (b) Drawing of dorsal view of caecum of midgut and pylorus, arranged laterally to the caecum of 
midgut, showing four branches and two pairs of tubular lateral diverticula, and (c) Drawing of dorsal view of 
brain and cerebral sensory organ (right side) showing relationships of these structure. Scale = 200 ym (b) and 
0.5 mm (c). ap, obliquely oriented projection of columnar portion of basis; bg, cyanophilous glandular mass of 
saclike canal; bm, branch of midgut caecum; cc, cover of central stylet; ci, collar-like ridge of massive posterior 


VOLUME 114, NUMBER 4 


glion cell on the left side was not observed. 
In paratype 1, large ganglion cells, 20 wm 
long and containing nuclei measuring 10 
jzm, lie medially in the brain, in the region 
of the ventral commissure (left) or behind 
it (right). It appears that the ganglion cells 
in this species are identical to Biirger type- 
3 ganglion cells, which are found in several 
cratenemertids (Biirger 1895). In the het- 
eronemertean Micrura leidyi, Riser (1998) 
demonstrated a neurochord cell in the gan- 
glionic layer of each ventral lobe of the 
brain and additional neurochord cells in the 
lateral nerve cords. His photomicrograph 
shows features similar to those of the large 
ganglion cell of N. fernaldi, but neuro- 
chords are lacking in both the brain and lat- 
eral nerve cords of this species. 

In the intestinal region, the lateral nerve 
cords, ventrolateral in position, lie above 
the longitudinal musculature of the body 
wall, and they give off dorsal or dorsoven- 
tral peripheral nerves that extend toward the 
dorsal and ventral sides of the body (Fig. 
8d). 

Several slender but conspicuous nerves 
originate from the brain (Fig. 6c). They 
branch out pre- and post-cerebrally to sup- 
ply various structures. A proboscis nerve 
trunk arises from the anterior side of each 
ventral ganglion and immediately extends 
to the proboscis sheath. Both proboscis 
nerves soon branch into seven nerves that 
lie on the lateral side of the sheath. Three 
small cephalic nerves, arranged one above 
the other and parallel to the proboscis nerve 
trunks, emerge from the anterior face of 
each ventral ganglion and proceed anteri- 


— 


847 


orly on the ventral side of the cephalic re- 
gion. Six small cephalic nerves originate 
from the dorsal ganglion a little farther pos- 
teriorly than the cephalic nerves on the ven- 
tral side, and they extend precerebrally to 
supply the eyes. A small nerve from each 
ventral ganglion extends anteriorly and then 
turns toward the outer side of the head, en- 
tering the dermis after passing the ventral 
side of the cerebral organ canal. A conspic- 
uous nerve originates from the lateral sur- 
face of each ventral ganglion and runs an- 
teriorly; its branches supply the oblique ce- 
phalic grooves. A small nerve from the lat- 
eral side of each dorsal ganglion extends 
anteriorly for a short distance and turns to- 
ward the lateral side of the head, innervat- 
ing the ocelli of the most posterior group. 

The midgut nerves originating from the 
medial side of each ventral ganglion run be- 
tween the ventral ganglia and the foregut, 
extending posteriorly along the dorsolateral 
portion of the foregut (Figs. 6c, 8e). The 
foregut nerves are a conspicuous feature of 
this species. They emerge on the ventral 
wall of the ventral ganglion, continue along 
the lateral side of the foregut, and have a 
commissure under the esophagus (Fig. 8e). 
The esophagus nerves, under the rhyncho- 
deum, emanate from the ventral surface of 
the ventral ganglia and extend anteriorly 
along the lateral sides of the esophagus, 
reaching beyond the brain. 

Three nerves, originating from the ven- 
tral side of each ventral ganglion, separately 
control different regions of the gut. One 
runs beneath the rhynchodeum and the an- 
terior part of the esophagus; one innervates 


portion of basis (the ridge encircles the columnar portion of the basis); cn, cerebral organ canal; co, conical part 
of posterior portion of basis, free of surrounding tissue; cp, columnar portion of basis; cs, central stylet; ct, 
bulbous part of posterior portion of basis, tightly bound to tissue; cu, caecum of midgut; dc, dorsal commissure 
of brain; dg, dorsal ganglion; er, esophagus nerve; fl, first pair of lateral diverticula of midgut caecum; fn, foregut 
nerve; gc, ganglionic mass of cerebral ganglion; gm, eosinophilous glandular mass of sensory canal; mi, midgut; 
mm, commissure of foregut nerve; mn, midgut nerve; nd, nerve to dermis; no, nerve to ocelli; ns, narrow duct 
of saclike canal; oc, opening of cerebral organ; op, opening of pylorus; ov, ovoid anterior portion of basis; pn, 
proboscis nerve; py, pylorus; sa, sensory canal; sl, secondary pair of lateral diverticula of midgut-caecum; ss, 
sac-type canal; tp, thick nerve from dorsal ganglion; vc, ventral commissure of brain; vg, ventral ganglion. 


848 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Seis Panay 


Fig. 7. Nipponnemertes fernaldi, new species. Transverse section of probosis apparatus in the holotype 
specimen (a), showing narrow canal in thick connective tissue of stylet bulb. Transverse sections of anterior 
portion of body, showing (b) mouth opening to rhynchodeum, (c) diverticulum of stomach, (d) pylorus, (e) 
vascular plug inside proboscis sheath, (f) nephridium, (g) thick nerve from dorsal ganglion to cerebral sensory 
organ, and (h) neurilemma and fibrous connective tissue of brain. Scale = 0.5 mm (a, c, d, h), 200 pm (6, f, 
g), 100 ym (e) and 50 pm (h). 37, number of section of proboscis apparatus (a) and 38—315; numbers of sections 
from anterior end of body (b—h). dg, dorsal ganglion; ds, diverticulum of stomach; ft, fibrous connective tissue; 
In, lateral nerve; lv, lateral blood vessel; mo, mouth; na, nephridium; nm, neurilemma; np, narrow canal in thick 
connective tissue and muscular wall of stylet bulb; ns, narrow duct of sac-type canal; pw, proboscis sheath, 
consisting of interwoven circular and longitudinal muscles; tp, thick nerve from dorsal ganglion; vg, ventral 
ganglion; vp, vascular plug inside proboscis sheath. 


VOLUME 114, NUMBER 4 849 


Fig. 8. Nipponnemertes fernaldi, new species. Transverse sections of anterior portion of body, showing (a, 
b) sensory canal and saclike canal after separation from cerebral organ canal, (c) large ganglion cell, (d) lateral 
nerve cord, (e) nephridium, (f) ocelli, (g) submuscular gland, and (h) posterior end of cerebral organ canal. Scale 
= 200 pm (a, b, e, g), 100 wm (d, f, h) and 50 wm (c). 129-188, numbers of sections from anterior end of 
body (a—c, e—h); 222, number of section from anterior end of middle portion of body (d). bt, beginning of T- 
shaped canal; cb, cellular body in dorsal ganglion; fn, foregut nerve; gc, ganglionic mass of cerebral sensory 
organ; mf, myofibril; mm, commissure of foregut nerve; mn, midgut nerve; ne large ganglion cell; ol, ocelli; 
sa, sensory canal; sg, shallow lateral groove; sm, submuscular gland; ss, sac-like canal; ts, sac epithelium; uc, 
U-shaped ciliated sensory portion; vp, vascular plug inside proboscis sheath. 


850 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


a 

Fig. 9. Nipponnemertes fernaldi, new species. Transverse sections of posterior portion of cerebral sensory 
organ. Nipponnemertes bimaculatus, transverse sections showing (c) sensory canal and sac (lumen T-shaped) 
and (d) sensory canal and sac without epithelium. Nipponnemertes punctatulus, transverse sections showing (e) 
sensory canal and sac (lumen T-shaped) and (f) sensory canal and sac without epithelium. Scale = 200 ym (d— 
f) and 50 pm (a, b). 173 and 176, numbers of sections from anterior end of body; 17, 20 and 22, 30, numbers 


of sections of proboscis apparatus (c—f). cy, cyanophilous glandular mass; dg, dorsal ganglion; eo, eosinophilous 
glandular mass; In, lateral nerve; ns, narrow duct of sac-like canal; sa, sensory canal; se, sac; si, sensory canal 


in eosinophilous glandular mass; ts, sac epithelium with T-shaped lumen. 


the posterior part of the esophagus, as well 
as the stomach and pylorus; the last inner- 
vates the midgut. 

The middorsal nerve lies in the dermis 
just above the circular muscle layer of the 
body wall and extends beyond the brain an- 
teriorly, below the dorsomedial longitudinal 
grooves (Fig. 2d). Its origin from the dorsal 
commissure of the brain was observed in 
paratype 1 but was not clear in the holo- 
type. The lateral nerves, with myofibrillae, 


give off peripheral nerves and are posteri- 
orly connected to each other by a commis- 
sure on the dorsal side of the rectum (Fig. 
8d). Myofibrillae in the lateral nerves are 
situated on the dorsal side between the out- 
er ganglion mass and the inner fibrous core. 
Myofibrillae have been described for some 
members of the Cratenemertidae (Crandall 
1993a, b; Crandall & Gibson 1998; Riser 
1998). 

Special sensory organs and frontal or- 


VOLUME 114, NUMBER 4 


gan.—In the holotype, the ocelli, counted 
in transverse sections of the head, number 
76. They are large and confined to the area 
between the flattened anterior end of the 
head and the middle region of the brain. 
There are 34 ocelli on the right side of the 
head and 35 on the left. Two groups on 
each side are recognizable; one consists of 
23 (right) and 24 (left) ocelli lateral to the 
rhynchodeum, and the second consists of 
11 (right and left) ocelli; three (right) and 
four (left) near the middle region of the 
brain are arranged dorsoventrally along the 
lateral sides of the head (Fig. 8f). 

In paratype 1, there are 78 ocelli, 39 on 
both sides. Of the two groups on the right 
side; one consists of 22 ocelli, and the other 
consists of 14; of the two groups on the left 
side, one consists of 25 ocelli, the other 
consists of 11. Three ocelli (right and left) 
near the middle region of the brain are ar- 
ranged as in the holotype specimen. In 
paratype 3 (Fig. lh), there are 52 ocelli, 26 
on both sides. 13 are marginal, 10 are on 
the anterior oblique cephalic grooves, and 
3 are behind it. The arrangement of ocelli 
in paratype 2 was not studied. 

The ocelli are of the inverted pigment- 
cup type and have dimensions of up to 90 
yum wide and 110 pm high (Fig. 8f). 

The cluster of frontal gland cells, which 
_ fan out into separate strands in the dorsal 
side of the head, is not large. In both the 
holotype and paratype 1 the cluster extends 
to the region of the proboscis insertion 
without reaching posteriorly to the cerebral 
ganglia (Figs. 3a—c, 7b). The frontal glands 
deliver their secretions to a pit at the ante- 
rior tip of the snout (Fig. 2g, h). The pit 
represents a frontal organ. 

The cephalic glands, confined to the lat- 
eral sides of the head, are found close to 
the frontal glands. The lateral bands of mu- 
coid glands are posterior extensions of the 
cephalic glands. Below the frontal organ, 
there is a narrow and short ciliated groove 
running posteriorly to the dorsal side of the 
rhynchodeal opening (Fig. 2g, h). In para- 


851 


type 1, this groove continues to the opening 
of the rhynchodeum. 

The posterior extensions of the mucoid 
cephalic glands, namely the submuscular 
glands, are situated laterally in the foregut 
region, beginning at the level of the anterior 
end of the cerebral organs (Fig. 8g). 

The cerebral organs, 510 wm long, are 
very large and extend behind the cerebral 
ganglia (Fig. 6c). For about 180 pm of its 
length, each is closely adjacent to the brain, 
which has a length of 460 wm. Anteriorly, 
the organ is circular in transverse section, 
but it becomes oval farther posteriorly (Fig. 
7g), and its posterior glandular portion is 
circular (Fig. 9b). The cerebral organ canal, 
ciliated and 220 wm long, begins at one of 
the secondary grooves of the anterior 
oblique cephalic grooves on the midlateral 
portion of the head, and is directed poster- 
omedially. It widens just before entering the 
cerebral organ and forms a U-shaped, cili- 
ated sensory portion (Fig. 8h). The epithe- 
lium of this is 20 wm thick. The canal has 
a widened lateral portion with a dorsally di- 
rected small groove lined with unciliated 
epithelium 8 pm thick (Fig. 8h). Farther 
posteriorly, the canal divides into a narrow 
sensory canal and a short, sac-like canal 
that has a T-shaped configuration when seen 
in transverse sections (Fig. 8a), and the dor- 
sally directed groove disappears (Fig. 8a, 
b). 

The sensory canal, 20 wm thick, lies on 
the side closest to the brain (Figs. 7g, 8a, 
b). It consists of a ciliated wall in which the 
sensory cells containing eosinophilous cy- 
toplasm are arranged in a row. Farther pos- 
teriorly, the ciliation disappears and the ca- 
nal enters a mass of eosinophilous glands 
about 40 wm long and 20 pm wide (Figs. 
6c, 9a, b). The total length of the canal is 
about 260 wm. 

The anterior portion of the sac-like canal 
(Fig. 6c), 80 wm by 60 pm in diameter and 
about 110 wm long, has a wall 10 wm thick. 
Its unciliated epithelium consists of slightly 
eosinophilic cells 6 4m wide; these are ar- 
ranged in one or two layers (Fig. 8b). Its 


852 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Summary of the main morphological characters of eleven marine nemerteans related to the genus 
Nipponnemertes including N. fernaldi, new species. 


Character Character states Coding 

1. Body color With pattern 0) 
Without pattern 1 

2. Cephalic grooves Two pairs of oblique cephalic grooves and dorsomedial longitudinal 0) 

grooves 

Two pairs of fluted cephalic grooves and inconspicuous oblique grooves 1 

A pair of shallow oblique fluted grooves 2 

A pair of lateral transverse grooves 3 

No record 4 

3. Cephalic grooves Anterior oblique cephalic groove subdivided by 14 secondary grooves, 0 


ventrolateral surface of head with 7 secondary grooves, posterior 
oblique cephalic grooves originating at the end of dorsomedial longitu- 
dinal grooves, and dorsomedial longitudinal grooves forming anteriorly 
a pair of epidermal folds 


No record 1 
4. Ocelli Numerous large ocelli in four groups 0 
In lateral rows 1 
5. Body wall Longitudinal muscle layer well developed 0) 
No record 1 
6. Diagonal muscles Present between circular and longitudinal muscle layers 0) 
No record 1 
7. Dermis Thick 0 
No record 1 
8. Rhynchocoel Nearly equal to body length 0) 
No record 1 
9. Proboscis sheath Composed of interwoven circular and longitudinal muscles 0 
No record 1 
10. Rhynchodeum Provided with a circular muscle layer 0) 
No record 1 
11. Precerebral septum Composed of muscular septum surrounding rhynchodeum 0 
Composed of muscular bundle 1 
No record 2 
12. Proboscis diaphragm Provided with central stylet-basis complex 0) 
Provided with a single central stylet and a barrel-, bell-, drop-shaped or 1 


conical basis 
No record 
13 Basis of central armature With posteriorly-inserted accessory stylet 
No record 
14. Stylet bulb With a plate-like muscular wall and a thick layer of glandular cells 
No record 
15. Accessory stylet pouches 2 
2 or more 
No record 
16. Proboscis nerves 11 
12-14 
14 
14-16 
15 
20 
No record 
17. Mouth Opening into rhynchodeum 
18. Esophagus Anterior to brain surrounded by longitudinal muscle 
No record 
19. Foregut caecum Esophageal caecum present 
Stomach caecum present 
No record 


NFP Or OCoaunBRWNRFPONK OF OF OW 


VOLUME 114, NUMBER 4 853 
Table 1.—Continued. 
Character Character states Coding 
20. Intestinal caecum Without anterior diverticula but with lateral ones 0 
With a pair of anterior diverticula sending off numerous branches 1 
No record 2 
21. Intestinal caecum and Intestinal caecum with anterior branches or pouches and two pairs of lat- 0) 
diverticula eral diverticula, and intestinal diverticula branched 
No record it 
22. Blood vascular system With three longitudinal vessels, middorsal vessel with single vascular 0 
plug 
No record 1 
23. Nervous system Brain and lateral nerves covered by a thin neurilemma and a thick layer 0 
of fibrous connective tissue, nervous system without neurochord cells, 
and accessory lateral nerves, middorsal nerve extending anteriorly be- 
yond brain, and lateral nerve cords with myofibrillae and peripheral 
nerves 
No record 1 
24. Ventral ganglia Not distinctly separated from dorsal ganglia 0) 
Posteriorly separated 1 
No record 2 
25. Dorsal ganglia Without bifurcated fiber core 0 
No record 1 
26. Nerve cords Paired midgut, foregut, and esophagus nerves originating from ventral 0) 
brain 
No record 1 
27. Frontal organ Present 0 
No record 1 
28. Cephalic glands Well developed and reaching posteriorly to cerebral ganglia 0) 
Prominent but reaching posteriorly to cerebral ganglia 1 
Moderately developed 2 
Small 3 
29. Submuscular glands Abundant in brain region 0 
Limited to a narrow area 1 
Absent 2 
No record 3 
30. Cerebral sensory organ lJLarge and extending behind cerebral ganglia 0) 
31. Cerebral sensory organ With neurilemma and cerebral organ canal branching into medial sensory 0) 
canal and lateral sac-like canal with two kinds of glandular masses 
No record 1 
32. Cerebral organ canal Open laterally in front of brain 0) 
Far anterior to brain 1 
No record 2 
33. Excretory system Pair of excretory pores lateral to nerve cords and situated in posterior 0 
portion of nephridial region 
Situated at middle of nephridial region 1 
Situated behind cerebral sensory organs D; 
No record 3 
34. Sexes Separated 0) 
No record 1 


very first part is still T-shaped in transverse 
section, and still has a shallow, dorsally di- 
rected lateral groove (Fig. 8a). The sac-like 
canal is succeeded by a narrow duct, 180 
wm long, surrounded by cyanophilous 


glands about 20 wm long and 15 wm wide 
(Figs. 6c, 7g, Qa). 

The internal configuration of the sensory 
canal and the sac-like canal can be under- 
stood by studying transverse sections at 


854 


their proper levels in Figures 6c, 7g, 8a, b, 
9a, b. The term “‘sac-like canal” is used 
here because this structure has a thick epi- 
thelium and also because it has cyanophi- 
lous glands (cf. Crandall 1993a). 

The ganglionic mass of the cerebral or- 
gan is mostly confined to the anterior por- 
tion of the organ, where it surrounds the 
sensory canal and sac-like canal. Glands are 
not present at the bifurcation of the cerebral 
organ canal (Fig. 8h). All of the organs are 
covered with connective tissue from the 
neurilemma of the cerebral nerve. The thick 
nerve from the posterior end of the dorsal 
ganglion enters the posterior portion of the 
glandular mass of the sac-like canal (Figs. 
6c, 7g). 

Excretory and reproductive systems.— 
The excretory tubules wind around and 
along the lateral blood vessels (Figs. 4a, 7c, 
d, f). In the holotype, they extend from the 
level of the posterior portions of the cere- 
bral sensory organs to the level of the pos- 
terior part of the pylorus; the region they 
occupy is 3.6 mm long. The efferent duct, 
beginning in the middle portion of the ne- 
phridial region, is conspicuous and extends 
above and lateral to the lateral nerve cord, 
reaching the excretory pore on the latero- 
ventral side of the body. The pore is ventral 
to the lateral nerve cord, and is at the level 
of the mid-pyloric region. The efferent duct 
of the left side, about 120 pm long, is much 
longer than that of the right side. The left 
excretory pore is located about 810 ym far- 
ther posterior than the right pore. 

In paratype 1, the excretory tubules ex- 
tend from the level of the posterior end of 
the cerebral sensory organs to a short dis- 
tance behind the pylorus; the region they 
occupy is 5.4 mm long. The efferent ducts, 
beginning in the posterior portion of the ne- 
phridial region, are 200 wm long; the ex- 
cretory pores are situated at their anterior 
ends. 

The ovaries of the fully mature holotype 
specimen are located near the beginning of 
the intestinal caecum, and are arranged ir- 
regularly between the intestinal diverticula 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(Figs. 2f, 4a, b). Gonoducts open at gono- 
pores on the lateral sides of the body. In the 
other two sectioned paratypes, both males, 
the gonads are undifferentiated and lack 
gonoducts. 

Swimming.—The specimens were ob- 
served to swim vigorously in the laboratory 
when the container was shaken. During 
swimming the body became more flattened 
and the swimming pattern was anguilli- 
form, resembling that of a Cerebratulus. 
Swimming has been described for the ho- 
plonemerteans Nipponnemertes punctatulus 
(Coe 1905) and N. pulcher (McIntosh 1873, 
Brunberg 1964, Berg 1972). 


Discussion of Systematics* 


The species described here is a member 
of the suborder Monostilifera (sensu Gibson 
1995), owing to the morphology of the ar- 
mature of the proboscis and the location of 
the mouth opening. It fits in the family Cra- 
tenemertidae (Friedrich, 1968) with respect 
to the following characteristics: rhyncho- 
coel wall with interwoven muscle layers of 
the proboscidial sheath; rhynchocoel ex- 
tending nearly to the posterior tip of the 
body; cerebral organs large, extending be- 
hind the cerebral ganglia (Berg 1985, Gib- 
son & Crandall 1989, Crandall & Gibson 
1998). 

The cratenemertid genus Nipponnemertes 
(type species: Nipponnemertes punctatulus 
(Coe, 1905), recently designated by Cran- 
dall (2001), is closest to the present species, 
especially with regard to the features cited 
in the generic diagnosis. Comparisons with 
other related species are found in Tables 1 
and 2. 

The arrangement of ocelli in N. fernaldi 
is virtually the same as that of all other Pa- 
cific Nipponnemertes, and the close juxta- 
position of the end of the rhynchocoel and 
the two posterior anastomoses also occurs 
in a few species of Nipponnemertes. In 
most cratenemertids, the anastomoses of 
blood vessels and lateral nerves are decid- 


VOLUME 114, NUMBER 4 


Table 2.—Character matrix for species of Nipponnemertes. Character numbers refer to Table 1. 


25 26) 9277/28) 29) BS 0 eS S23) 64: 


24 


13} 4h IS) th Gf ile} 19 2022228) 


12 


11 


10 


Character 


2 6 O 
0 2 0 


0 


N. drepanophoroides 


N. pacificus 


2 
2 2 (— 
0 0 O 


0 0 0 


1 


0 6 O 


N. occidentalis 
N. punctatulus 


1 
1 


0 0 O 


0 0 O 


N. bimaculatus 
N. pulcher 


0 2 0 
0 2 0 


1 


0 0 0 O 


0 0 O 
ee ko) 
0 4 0 
0 2 O 


0 0 O 


N. africanus 


N. magnus 
N. marioni 


0 0 3 0 


1 


0 O 


0 0 O 


1 


0 0 0 0 O 


0 O 


0 
0200000 0 0 0 OF! 


N. schollaerti 
N. fernaldi 


it 


© @O @O ®@ Oi! @® O O 2 


2) 


1 


0) 


855 


edly posterior to the end of the rhyncho- 
coel. 

Concerning the structure of the cerebral 
organs, Gibson (1988) stated that the cere- 
bral organs of Antarctic species of Nippon- 
nemertes resemble those of inaequifurcate 
Eureptantia in being large and elaborate 
structures, close to and reaching behind the 
cerebral ganglia; furthermore, their cerebral 
canals are divided. Gibson (1988) also 
pointed out that the cerebral organs of Nip- 
ponnemertes appear much more highly 
evolved than those of almost all Monostil- 
ifera, and that their organization and posi- 
tion may be indicative of a closer evolu- 
tionary relationship to Reptantia than has 
previously been appreciated. On the basis 
of these features, Gibson (1988) pointed out 
that there are similarities between Nippon- 
nemertes and Reptantia (Inaequifurcata). 
Crandall (1993a) wrote that N. bimaculatus 
(Amphiporus bimaculatus), now known to 
be a cratenemertid, possesses a canal that 
divides, just after entering the cerebral or- 
gan, into two branches: a sensory branch 
having a uniform elliptical cross-section 
throughout its length, and a shorter branch 
of similar size but complex shape, lined 
with characteristic ‘sac-like’ epithelium. 

Nipponnemertes bimaculatus from Rishi- 
ri Island in Hokkaido, Japan (Iwata 1954) 
has large cerebral organs situated at the lat- 
eral side of the brain and extending poste- 
riorly behind the brain. In addition, the 
muscle layer of the proboscis sheath is in- 
terwoven, a feature clearly evident in my 
slides. The cerebral organs are like those in 
the present species (Fig. 9c, d). Nipponne- 
mertes punctatulus from Onomichi (Iwata 
1951) has several characters of the genus: 
large cerebral organs situated at the lateral 
side of the brain and extending posteriorly 
behind the brain; the interwoven muscle 
layer of the proboscis sheath; similarity of 
the cerebral organs to those of N. bimacu- 
latus (Fig. 9e, f); and a distinct rectum with 
circular muscle sphincter. 

Nipponnemertes fernaldi is distinctive 
because of the following features: a central 


856 


stylet-basis complex of unique structure; a 
bulblike stylet bulb with a platelike mus- 
cular wall and a thick layer of glandular 
cells; an intestinal caecum with four 
branches or pouches and two pairs of lateral 
diverticula; a pair of dorsomedial longitu- 
dinal epidermal folds in the anterior region. 
These features have not been reported for 
cratenemertids of the genus Nipponnemer- 
tes until now. The combination of charac- 
ters found in N. fernaldi decidedly excludes 
it from the existing species of Nipponne- 
mertes (Griffin 1898, Coe 1901, 1905; Pun- 
nett 1903, Wheeler 1934, 1940; Iwata 1951, 
1954; Friedrich 1968, Berg 1972, 1985; 
Gibson 1995, Crandall & Gibson 1998). 
Because of this, a new species is proposed 
to accommodate it. 


Acknowledgments 


I thank A. O. Dennis Willows, Director 
of Friday Harbor Laboratories, University 
of Washington, for his constructive com- 
ments and for providing facilities for re- 
search. Eugene Kozloff, also of Friday Har- 
bor Laboratories, helped in preparation of 
the manuscript. Finally, Jon Norenburg, and 
Frank B. Crandall of the National Museum 
of Natural History, Washington, D. C. and 
Stephen L. Gardiner of Bryn Mawr College 
gave me useful comments and suggestions. 
The help of an anonymous reviewer is also 
appreciated. This study was supported in 
part by a NSF Grant, USA in 1964 and by 
the Kushiro Public University of Econom- 
KES iim WYYS). 


Literature Cited 


Berg, G. 1972. Studies on Nipponnemertes Friedrich, 
1968 (Nemertini, Hoplonemertini). I. Rede- 
scription of Nipponnemertes pulcher (Johnston, 
1873) with special reference to intraspecific var- 
lation of the taxonomic characters.—Zoologica 
Scripta 1:211—225. 

. 1985. Studies on Nipponnemertes Friedrich 
(Nemertini, Hoplonemertini). Il. Taxonomy of 
Nipponnemertes pulcher (Johnston) and some 
other species.—Zoologica Scripta 14:239—246. 

Brunberg, L. 1964. On the nemertean fauna of Danish 
waters.—Ophelia 1:77—111. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Birger, O. 1895. Die Nemertinen des Golfes von Nea- 
pel und der angrenzendes Meeres-Abschnitte.— 
Fauna und Flora des Golfes von Neapel 22:1— 
743. 

Coe, W. R. 1901. Papers from the Harriman Alaska 
Expedition. XX. The nemerteans.—Proceedings 
of the Washington Academy of Sciences 3:1— 
110. 

. 1905. Nemerteans of the west and north-west 
coasts of America.—Bulletin of the Museum of 
Comparative Zoology at Harvard College 47:1— 
318. 

Crandall, KE B. 1993a. Major characters and enoplan 
systematics.—Hydrobiologia 266:115—140. 

. 1993b. The genus Valdivianemertes Stiasny- 

Winhoff 1923 (Nemertea, Enopla, Hoplone- 

mertea): nomenclatural status and proper sys- 

tematic position.—Hydrobiologia 266:175—184. 

. 2001. A cladistic view of the Monostilifera 

(Hoplonemertea) with interwoven rhynchocoel 

musculature: a preliminary assessment.—Hy- 

drobiologia (in press). 

, & R. Gibson. 1998. A second genus of pelagic 
Cratenemertidae (Nemertea, Hoplonemertea).— 
Hydrobiologia 365:173—198. 

Friedrich, H. 1968. Sagaminemertes, eine bemerken- 
swerte neue Gattung der Hoplonemertinen und 
ihre systematische Stellung.—Zoologischer An- 
zeiger 180:33—36. 

Gibson, R. 1988. Evolutionary relationships between 


mono- and polystiliferous: Nipponnemertes 

(Cratenemertidae), a missing link genus?—Hy- 

drobiologia 156:61—74. 

. 1995. Nemertean genera and species of the 

world: an annotated check-list of original names 

and description citations, synonyms, current 
taxonomic station, habitats and recorded zoo- 
geographic distribution.—Journal of Natural 

History 29:271—562. 

, & FE B. Crandall. 1989. The genus Amphi- 
porus Ehrenberg, 1831 (Nemertea, Enopla, 
Monostiliferoidea).—Zoologica Scripta 18: 
453-470. 

Griffin, B. B. 1898. Description of some marine ne- 
merteans of Puget Sound and Alaska.—Annals 
of the New York Academy of Science 11:193— 
Dilip 

Iwata, EF 1951. Nemerteans in the vicinity of Onomi- 
chi.—Journal of the Faculty of Science, Hok- 
kaido University, Series 6, Zoology 10:135— 
138. 

. 1954. The fauna of Akkeshi Bay. XX. Nem- 

ertini in Hokkaido.—Journal of the Faculty of 

Science, Hokkaido University, Series 6, Zool- 

ogy 12:1—-39. 

. 1998. On the hoplonemertean, Kameginemer- 

tes parmiornatus (Iwata, 1957) gen. n., comb. 


VOLUME 114, NUMBER 4 


n. from Sagami Bay, Japan.—Hydrobiologia 
365:199—213. 

McIntosh, W. C. 1873--1874. A monograph of British 
Annelids. Part 1. The Nemerteans.—London: 
Ray Society: 1—244. 

Punnett, R. C. 1903. On the nemerteans of Norway.— 
Bergens museums arbog. 2:1—35. 

Riser, N. W. 1998. The morphology of Micrura leidyi 
(Verrill, 1892) with consequent systematic re- 
valuation.—Hydrobiologia 365:149—156. 

Stricker, S. A. 1985. The stylet apparatus of monostil- 
iferous hoplonemerteans.—American Zoologist 
25:87-97. 

, & R. A. Cloney. 1981. The stylet apparatus 

of the nemertean Paranemertes peregrina: its 


ultrastructure and role in prey capture.—Zoo- 
morphology 97:205—223. 

Wheeler, J. E G. 1934. Nemerteans from the South 
Atlantic and southern oceans.— ‘Discovery’ Re- 
ports 9:215—294. 

. 1940. Some nemerteans from South Africa 


857 


and a note on Lineus corrugatus McIntosh.— 
Journal of the Linnean Society 41:20—49. 


* Editor’s note: The author mentioned 
the taxon as Nipponnemertes fernaldi nov. 
sp. in an abstract published in the Japanese 
journal Taxa (no. 11, 2001) in September 
2001. However, this publication of it did 
not make the name ‘available’, since no 
type specimen was designated therein as re- 
quired by Article 72.3 of the International 
Code of Zoological Nomenclature (4th Ed., 
1999). The full description and designation 
of type specimens in this issue of the Pro- 
ceedings of the Biological Society of Wash- 
ington 114(4):833-857, make the name 
available for the first time, and it is cor- 
rectly used therein as Nipponnemertes fer- 
naldi n. sp. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(4):858—860. 2001. 


Dhondtichlamys, a new name for Microchlamys Sobetski, 1977 
(Mollusca: Bivalvia: Pectinidae), preoccupied by Microchlamys 
Cockerell, 1911 (Rhizopoda: Arcellinida) 


Thomas R. Waller 


Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, 
Washington, D. C. 20560-0121, U.S.A. 


Abstract.—The Cretaceous pectinid subgenus name Chlamys (Microchla- 
mys) Sobetski, 1977, is preoccupied by Microchlamys Cockerell, 1911, a pro- 
tozoan. The new name Dhondtichlamys is introduced to replace the junior 


homonym. 


Sobetski (1977:56) introduced a new 
subgenus, Chlamys (Microchlamys), for a 
group of Eurasian Upper Cretaceous bi- 
valves of the family Pectinidae. The type 
species designated by Sobetski is Pecten 
pulchellus Nilsson, 1827, the type stratum 
and locality of which are the Campanian 
greensands at K6pingemolla, Sweden 
(Dhondt 1972:19). These Cretaceous glau- 
conitic sandstones occur in the K6pinge 
district, which includes K6pingem6lla, and 
are now referred to by the informal name 
““K6pinge sandstone.”’ Christensen (1986) 
described two belemnite assemblages from 
it, the older indicating a latest Early Cam- 
panian age and the younger a middle Late 
Campanian age. Kennedy and Christensen 
(1997) recorded ammonites from this sand- 
stone that indicated an age compatible with 
the younger belemnite assemblage. Nils- 
son’s material likely came from that part of 
the K6pinge sandstone that yielded the am- 
monites and therefore is probably of middle 
Late Campanian age (W. K. Christensen, in 
litt.). 

Vokes (1980:229) reported that Sobet- 
ski’s subgenus name is preoccupied by Mi- 
crochlamys Cockerell, 1911 (a rhizopodan 
protozoan of the order Arcellinida), but So- 
betski did not rename the taxon before his 
death in 1988. As required by the Interna- 
tional Code of Zoological Nomenclature 


(Fourth Edition, Article 60.3), I replace Mi- 
crochlamys Sobetski, 1977, with Dhond- 
tichlamys, new name, based on the same 
type species, Pecten pulchellus (ICZN Rec- 
ommendation 60A). The name honors Dr. 
Annie V. Dhondt, Koninklijk Belgisch In- 
stituut voor Natuurwetenschappen, Brus- 
sels, for her many contributions to our 
knowledge of Cretaceous Pectinidae. She 
was the first (Dhondt 1972) to recognize the 
taxonomic significance of the group of spe- 
cies that Sobetski later named. 

In addition to the type species, Dhondt- 
ichlamys includes eight of the nine species 
that Sobetski (1977) included in Micro- 
chlamys: Pecten acuteplicatus Alth, 1850; 
P. arlesiensis Woods, 1902; P. campanien- 
sis d’Orbigny, 1847; P. inflexus Hagenow, 
1842; P. puggaardi Ravn, 1908; Chlamys 
(Microchlamys) subacutiformis Sobetski, 
1977; P. subaratus Nilsson, 1827; and 
Chlamys (Aequipecten) wisniowskii Paster- 
nak, 1961. He originally also included P. 
trisulcus Hagenow, 1842, but later (Sobet- 
ski in Sobetski et al. 1982) moved this spe- 
cies to Chlamys (Chlamys). Dhondt (1972) 
had earlier included essentially the same as- 
semblage, with the addition of Pecten sar- 
umensis Woods, 1902, under the now ob- 
solete combination “‘Lyropecten (Aequipec- 
ten).”’ She determined that P. inflexus is a 
junior homonym, which she renamed Lyr- 


VOLUME 114, NUMBER 4 


opecten (Aequipecten) subinflexus Dhondt, 
1972. She also regarded Pecten puggaardi 
and probably also P. wisniowskii as junior 
synonyms of Pecten campaniensis. Sobet- 
ski et al. (1982) added three new species, 
Chlamys (Microchlamys) acuticostata So- 
betski, 1982, C. (M.) plestshejevi Sobetski, 
1982, and C. (M.) entis Sobetski, 1982. 

Collectively, the above assemblage of 
species ranges in age from Albian to Maas- 
trichtian and occurs in Great Britain, Eur- 
asia, and Egypt. As noted by Dhondt (1972: 
13, 40), the genus also includes a species 
from the Upper Cretaceous of eastern North 
America, Pecten venustus Morton, 1833. 
Darragh & Kendrick (1991:55) described a 
new species, Chlamys (Microchlamys) pro- 
pesalebrosa, from the Maastrichtian of 
northwestern Australia, this being the first 
record of the genus in the Southern Hemi- 
sphere. Stilwell (1998:44), however, ques- 
tioned this generic placement. It is likely 
that Dhondtichlamys survived the end-Cre- 
taceous mass extinction (Waller 1991, Wal- 
ler & Marincovich 1992), but Paleocene or 
Eocene specimens remain poorly docu- 
mented (for example, Pecten farafrensis 
Zittel, 1883, a nomen nudum discussed by 
Dhondt 1972:28). Waller (1991) and Waller 
& Marincovich (1992) used Microchlamys 
at the genus rather than the subgenus level 
for phylogenetic reasons, a usage that was 
followed by Dhondt et al. (1996:57) and 
Dhondt & Jagt (1997:44). 


Acknowledgments 


I thank W. K. Christensen, A. V. Dhondt, 
D. Jablonski, L. Marincovich, Jr., and A. E. 
Oleinik for useful information and/or com- 
ments on the manuscript. 


Literature Cited 


Alth, A. 1850. Geognostisch-palaontologische Bes- 
chreibung der naéchsten Umgebung von Lem- 
berg.—Naturwissenschaftliche Abhandlungen, 
gesammelt und durch subscription herausgeben 
von Wilhelm Haidinger, Abhandlungen 3:171— 
284. 

Christensen, W. K. 1986. Upper Cretaceous belemnites 


859 


from the Vomb Trough in Scania, Sweden.— 
Sveriges Geologiska Unders6kning Ca57:1—57. 

Cockerell, T. D. A. 1911. The nomenclature of the Rhi- 
zopoda.—Zoologischer Anzeiger 38:136—137. 

Darragh, T. A., & G. W. Kendrick. 1991. Maastrichtian 
Bivalvia (excluding Inoceramidae) from the 
Miria Formation, Carnarvon Basin, north west- 
ern Australia.—Records of the Western Austra- 
lian Museum, Supplement No. 36:1—102. 

Dhondt, A. V. 1972. Systematic revision of the Chla- 
mydinae (Pectinidae, Bivalvia, Mollusca) of the 
European Cretaceous. Part 2: Lyropecten.— 
Bulletin Instituut royal des Sciences naturelles 
de Belgique 48(7):1—81. 

, & J. W. M. Jagt. 1997. Late Cretaceous 

macrofauna from the Hautes Fagnes area (NE 

Belgium).—Annals de la Société géologique de 

Belgique 119(1):39—53. 

, D. P. Naidin, A. P. Levina, & E. Simon. 1996. 
Maastrichtian faunas from the Turgay Strait 
(northern Kazakhstan).—Mitteilungen aus dem 
Geologisch-Paldontologischen Institut der 
Universitat Hamburg 77:49-65. 

Hagenow, E von. 1842. Monographie der Rtigen’schen 
Kreidevertseinerungen III. Mollusken.—Neues 
Jahrbuch fiir Mineralogie, Geologie und Paléon- 
tologie 1842:528—575. 

International Commission on Zoological Nomencla- 
ture, 1999. International Code of Zoological 
Nomenclature, Fourth Edition. The Internation- 
al Trust for Zoological Nomenclature c/o The 
Natural History Museum, London, 306 pp. 

Kennedy, W. J., & W. K. Christensen. 1997. Santonian 
to Maastrichtian ammonites from Scania, south- 
ern Sweden.—Fossils and Strata 44:75—-128. 

Morton, S. G. 1833. Supplement to the “Synopsis of 
the organic Remains of the Ferruginous Sand 
Formation of the United States,’ contained in 
Vols. XVII and XVIII of this Journal.—Amer- 
ica Journal of Science and Arts 23(2):288—294. 

Nilsson, S. 1827. Petrificata suecana formationis cre- 
taceae, descripta et iconibus illustrata. Pars Pri- 
or, Vertebrata et Mollusca sistens. Londini 
Gothorum, Lund, Sweden, 39 pp. 

Orbigny, A. d’. 1847. Paléontologie frangais—De- 
scription des Mollusques et Rayonnés fossi- 
les—Terrains crétacés. IlJ. Lamellibranches. G. 
Masson, Paris, pp. 521—807. 

Pasternak, S. I. 1961. On the systematics of Upper 
Cretaceous pectinids.—Paleontologischeskii 
Sbornik 1:19—21. 

Ravn, J. P. J. 1908. Fortgn. over Kridtforst. fra Stevns 
Klint—Danmarks geologiske. undersggelse, 
Series I, 11:66—76. 

Sobetski, V. A. 1977. [Bivalve molluscs of the Late 
Cretaceous platform seas of southwestern 
USSR.]—Trudy paleontologicheskogo Instituta 
Akademii Nauk SSSR 159:1—256 [in Russian]. 


860 


, et al. 1982. [An atlas of invertebrates of Late 
Cretaceous seas of the Caspian Syncline.]— 
Trudy Paleontologischeskogo Instituta Akade- 
miya Nauk SSSR, Moscow, 187:1—252 [in Rus- 
sian]. 

Stilwell, J. D. 1998. Late Cretaceous Mollusca from 
the Chatham Islands, New Zealand.—Alcher- 
inga 22:29-85. 

Vokes, H. E. 1980. Genera of the Bivalvia: A system- 
atic and bibliographic catalogue (revised and 
updated). Paleontological Research Institution, 
Ithaca, New York, 307 pp. 

Waller, T. R. 1991. Evolutionary relationships among 
commercial scallops (Mollusca: Bivalvia: Pec- 
tinidae). Pp. 1-73 in S. E. Shumway, ed., Scal- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


lops: Biology, Ecology and Aquaculture. Elsev- 

ier Science Publishers, New York, 1095 pp. 

, & L. Marincovich, Jr. 1992. New species of 
Camptochlamys and Chlamys (Mollusca: Biv- 
alvia: Pectinidae) from near the Cretaceous/Ter- 
tiary boundary at Ocean Point, North Slope, 
Alaska.—Journal of Paleontology 66:215—227. 

Woods, H., 1902. A monography of the Cretaceous 
Lamellibranchia of England.—Palaeontograph- 
ical Society Monograph, Vol. I, Pectinidae: 
145-232. 

Zittel, K. A. 1883. Beitrage zur Geologie und Palaon- 
tologie der libyschen Wiiste und der angrenzen- 
den Gebiete von Aegypten. I. Geologischer 
Theil.—Palaeontolographica 30(1):1—147. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
114(4):861—880. 2001. 


Sipuncula from Antarctic waters 


Edward B. Cutler, Harlan K. Dean, and José I. Saiz-Salinas 


(EBC, HKD) Department of Invertebrate Zoology, Museum of Comparative Zoology, 
26 Oxford St, Cambridge, Massachusetts, 02138, U.S.A.; 


(JIS-S) Dpto. de Zoologia y DCA, Universidad Pais Vasco E-48080 Bilbao, Apdo. 644, Spain 


Abstract.—A collection of 3156 Sipuncula from water south of 53° S latitude 
collected by several American expeditions during 1948-1986, at depths from 
1—5790 m yielded 11 species. Only Apionsoma murinae (both subspecies) are 
new to this region but the known longitudinal ranges of the others are ex- 
panded. There is one numerically dominant species in each of the other three 
genera: Golfingia margaritacea, Phascolion lutense and Nephasoma diaphanes. 
Comments on the morphology of each taxon are presented plus a general ob- 
servation about the deciduous nature of introvert hooks across these four gen- 
era. An illustrated key and distribution maps are provided. 

Resumen.—Se estudia una colecci6n de 3156 ejemplares de sipunctlidos 
procedentes de los mares antdarticos situados por debajo de los 53° latitud sur, 
siendo encontrandas un total de 11 especies. Apionsoma murinae (ambas su- 
bespecies) representan citas nuevas para la zona investigada, mientras que se 
amplian los rangos de distribuci6n longitudinal de las otras especies identifi- 
cadas. Una de las especies de cada uno de los géneros restantes: Golfingia 
margaritacea, Phascolion lutense y Nephasoma diaphanes, siempre es partti- 
cularmente abundante en la colecci6n. Asimismo se presentan comentarios so- 
bre la morfologia de cada taxon identificado, destacandose la naturaleza caediza 
de los ganchos de la trompa en los 4 géneros investigados. Finalmente se 
incluyen claves de identificaci6n y mapas de distribucion de todas las especies 


tratadas. 


This report is based on 3156 specimens 
collected at 456 stations over 38 years 
(1948-1986) by several American research 
vessels, Operation Deepfreeze contributed a 
few in 1948, then no more until 1956. Most 
of the collecting was done over the next 
two decades, with less intensity from the 
mid 1970’s through the mid 1980’s. Among 
the ships making the largest contribution 
are the R/V Eltanin, Hero, Glacier, and Is- 
las Orcadas, but there were several others 
as well as a few land based operations. 

Initial processing was by the Smithsonian 
Oceanographic Sorting Center (SOSC). The 
samples came from depths of 1—5790 m and 
latitudes of 53—78.5° S. This is an arbitrary 


northern boundary since there are speci- 
mens scattered throughout the oceans of the 
world, including some from the Arctic 
Ocean. These will be treated in other re- 
ports. Several dozen Antarctic specimens 
were omitted from this report due to their 
damaged or incomplete nature. The nomen- 
clature used here follows that of Cutler 
(1994). 


Materials and Methods 


Specimens were collected using a wide 
range of trawls, dredges and grabs. The lo- 
cation and number of specimens are listed 
in the text if there were five or fewer sta- 
tions. For more commonly collected species 


862 


this is summarized in the text and the de- 
tailed data can be found in Appendix I. For 
identification, standard dissecting tech- 
niques and both binocular dissecting and 
compound microscopes were used as re- 
quired. The specimens are housed in the 
National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 


Results 


Key to the species in this report (see Fig. 
1D): 


1. Single nephridium ... (genus Phascolion) 
— Two nephridia 
2. Introvert retractor muscles appear as 
two separate units and of nearly equal 
SIZ, ae Phascolion (Isomya) hedraeum 
— Introvert retractor muscles fused into 
single column with very short distinct 
origins (subgenus Montuga) 
3. Trunk smooth, without apparent hold- 
fast papillae, lives in clay tubes 
Phascolion lutense 
— Trunk with thin but distinct holdfast pa- 
pillae P. pacificum 
4. One pair introvert retractor muscles .. 
(genus Nephasoma) 
— Two pair introvert retractor muscles .. 7 
5. Elongate very slender trunk, sometimes 
threadlike (width usually <0O.1 the 
length); gut with separated coils 
Be eeea obi, Seo Nephasoma capilleforme 
— Cylindrical trunk (width rarely <0.1 the 
length); gut coils close together 
6. Introvert hooks spine-like (one edge 
shorter than others) <50 pm tall 
Nephasoma diaphanes 
— Hooks pyramidal, >50 pm tall ..... 
ny aes Le Nephasoma abyssorum benhami 
7. Introvert more than twice length of 
trunk .... (Apionsoma murinae)..... 8 
— Introvert less than twice trunk length 
(Golfingia) 
32 Nephiidiarcimele-lobediin = ite eee 
Apionsoma murinae murinae 
— Nephridia with secondary lobe 


Apionsoma murinae bilobatae 
9. Posterior trunk without caudal append- 
AgeDA Tee Ve Golfingia margaritacea 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


— Posterior trunk with caudal appendage 


10. Large bladder-like papillae at base of 
(21 ble anf cer ere ots Golfingia anderssoni 
— Base of tail without large bladder-like 
Papilllacsy..o tay ee G. muricaudata 


Note: If one has worms less then 2 or 3 mm 
long, one must use more comprehensive sources 
to go beyond the generic level. 


Genus Golfingia Lankester, 1885 
Golfingia anderssoni (Théel, 1911) 


Material examined.—48 specimens from 
60 stations at latitudes 53—78° S, longitudes 
from 26—64° W, then skip to 160° W—165° 
E. The total depth range was 22-3413 m, 
but 70% of the samples from 100—1200 m 
(Fig. 5a). 

Description.—The trunks range from 2— 
133 mm in length; most from depths >1000 
m are >35 mm long, while those from shal- 
lower water are mostly <10 mm long. Un- 
like data published prior to Saiz (1995) (he 
had specimens up to 110 mm from the Ant- 
arctic) that records no specimen longer than 
35 mm, this collection has 44 worms (30%) 
that exceed this length. A surprising num- 
ber of specimens were unusually elongated 
and thin reflecting the highly plastic nature 
of this body and perhaps a distortion of 
their living state. 

The posterior end of most specimens ex- 
hibits the characteristic large, bladder-like 
papillae on the posterior quarter of the trunk 
and thin, rat-like caudal appendage (tail, 
Fig. 2a). In worms less than 8 mm long 
these attributes may be under-developed 
(Fig. 2b) and inspection under a compound 
microscope may be necessary to differen- 
tiate this from small G. muricaudata. Oc- 
casionally, these papillae resembled blad- 
ders that had been drained, thus looking 
more like flattened scales. Tails can be bro- 
ken during collection, but in a sample of 24 
worms 11—133 mm long, the tails are 4— 
16% of the trunk plus one each at 22 and 
33%. 

While earlier observers did not see intro- 


VOLUME 114, NUMBER 4 


vert hooks on this species, we observed 
small (30-35 wm), triangular, pale, scat- 
tered hooks on a number of specimens (Fig. 
2c). These could not be seen with the dis- 
secting microscope but required examina- 
tion under the compound scope. However, 
as predicted in Cutler (1994), these hooks 
are numerous on 2—6 mm worms. They are 
still present but few on 9-12 mm worms 
and absent in 17 and 30 mm worms from 
the same station. 

A single worm from the most southerly 
station (USNM 170138) displayed what we 
interpret to be an anomalous condition in 
that the dorsal pair of introvert retractor 
muscles is absent. If this worm were col- 
lected from another part of the world it 
would very likely be identified as Nepha- 
soma flagriferum. The occasional absence 
of one or two retractor muscles in Golfingia 
species has been commented previously 
(Gibbs 1973). 

Distribution.—Of the 14 previous reports 
between 1911—1996, all but three are from 
the far southern latitudes at depths of 75— 
1880 m. One is from the equatorial Atlantic 
Ocean (4 worms near 18° S, 9’ E) at 4300— 
4600 m. This might be a case of equatorial 
submergence or a mistake (Cutler & Cutler 
1987). Two northern Pacific Ocean reports 
of single worms (28 and 44° N) from deep 
water (3150 and 6135 m), are difficult to 
reconcile with what seems to be a southern 
ocean taxon (Murina 1964, 1971). The pre- 
sent data strengthens the concept of this as 
a circum-Antarctic bathyal species. 


Golfingia margaritacea (Sars, 1851) 


Material examined.—These 1442 speci- 
mens came from 305 stations at depths 
from 1—4886 m. However, 90% came from 
50—1200 m (9% <50, none between 1200— 
2273 and only 1% >2200 m). The full 
range of latitudes from 53—78.5° S is rep- 
resented but the longitudinal coverage is 
discontinuous. Going westward from 0°, 
there are scattered records from 16 and 22°, 
then the bulk from 26—70°, then scattered 


863 


again at 110—115°, 158° and 166° W, then 
another heavy concentration between 173° 
W and 159° E. A final few came from 150°, 
116-110°, 101°, and 93° E (Fig. 5b). 

Description.—This most common Ant- 
arctic species (45% of the material) ranges 
in trunk length from 1-79 mm. Those 
worms exceeding 25 mm lived at depths 
>150 m, mostly >300 m. One exception is 
the two intertidal worms from the tip of 
South America (USNM 170049) measuring 
23 and 34 mm. Small worms dominated 
this collection with 94% being <10 mm 
long, while 4% are 10—25 mm, and only 
2% are >25 mm in trunk length. 

While most specimens are cylindrical 
(ength 4—8 times width) with rounded or 
bluntly pointed conical posterior ends (oc- 
casionally with a dimple or small depres- 
sion), others are elongate and thin while 
some are short and fat. The color varies 
from pale through orange to dark brown 
and the skin which is usually thin and 
smooth can be quite thick and rugose. The 
more slender introvert in the smaller worms 
is about equal to the trunk in length, but 
due to allometric growth is only about half 
the trunk length in larger specimens. Also, 
in small worms (1.5—8 mm) small (20-35 
jum) scattered hooks exist that are lost as 
the worm matures. Papillae are more dense 
near the extremities of the trunk, and are 
generally small digitiform structures, but 
their size is remarkably variable. 

Internally, one can often observe vesicles 
or bubbles on the swollen contractile vessel. 
Two individuals had only 3 retractor mus- 
cles, one of the dorsals being absent, a phe- 
nomenon known in other populations (e.g., 
Gibbs 1973). Within the coelomic cavity of 
several worms maturing Ova were seen. 
These worms were 6—17 mm long and had 
been collected between December and May, 
most during February and March. 

Note.—When Cutler (1994:73) retained 
the sub-species G. margaritacae ohlini 
(Théel, 1911), it was with some hesitation 
since there were only two indistinct differ- 
ences, i.e. hooks sometimes present and a 


864 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Schematic drawings of the sipunculans to illustrate the identification key. Numbers are in accordance 
with the couplet numbers of the key. 


VOLUME 114, NUMBER 4 


Fig. 1. Continued. 


865 


Senn 
C Ne! 
— == 
Fig. 2. Small specimens (less than 8 mm trunk 


length) of Golfingia anderssoni. A: Variation in caudal 
appendage development. B: Bladder-like papillae. C: 
Hooks. Scale bars: A = 2 mm; B = 50 wm; C = 20 
pm. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


variably pointed posterior end. When Saiz 
(1995) examined a large collection of Wed- 
del Sea specimens, he concluded that any 
attempt to retain a distinction between these 
putative sub-species was an exercise in fu- 
tility. We affirm that conclusion. 
Distribution.—Very wide-spread in the 
Atlantic, Arctic, and Antarctic oceans (80° 
N to 78° S). In the Pacific Ocean at higher 
latitudes (>30° N & S) and on those oc- 
casions when reported from lower latitudes 
it is from depths >2000 m. Recent Indian 
Ocean records are limited to sub-Antarctic 
latitudes (46—50° S) most from near the 
Kerguelen Islands. The reported depth 
range is 1—5300 m, but most specimens 
have been collected from depths less than 
400 m. Eighty six percent of the specimens 
reported here were from that depth range, 
and while filling in some gaps in longitu- 
dinal distribution there are no surprises. 


Golfingia muricaudata (Southern, 1913) 


Material examined.—69 specimens from 
30 stations at latitudes 54—77.8° S, and the 
bulk between longitudes 27.5—65° W, plus 
a few at 166—170° E, and a single station at 
90.6° E. Depths range from 20-892 m, but 
80% of these stations were at 100—600 m 
(Fig. 5a). 

Description.—The trunks in this collec- 
tion range from 2—19 mm long, but only 5 
(7%) are over 10 mm long, thus somewhat 
smaller than elsewhere. The characteristic 
caudal appendage easily seen in larger 
worms, may be only a small pointed bump 
in worms less than 5 mm long. These small 
worms are not easy to differentiate from G. 
anderssoni except for the papillae on the 
posterior trunk which are very small, thin, 
and hair-like, not large bladders (Fig. 3a). 
Scattered hooks, known to be deciduous, 
were observed on a few small (4—7 mm) 
worms (Fig. 3b). 

Not included above are two worms from 
USNM 170708 that are much larger (32 & 
60 mm) and collected from much greater 
depths (2975 m). While such depth and size 


VOLUME 114, NUMBER 4 


867 


Fig. 3. 


Small specimens (less than 7 mm trunk length) of Golfingia muricaudata. A: Caudal appendage 


showing thin papillae. B: Hooks. Scale bars: A = 200 pm; B = 15 pm. 


are not unusual for the species they are very 
striking in this collection. Their caudal ap- 
pendages are 28 & 17%, respectively. Of 
particular interest is the fact that at this 
same station we found two examples of the 
very similar G. anderssoni of similar large 
Size. 

Distribution.—A widely distributed but 
low density cold water species in the higher 
latitudes of the Atlantic, Pacific, and Indian 
Oceans (e.g., Bering and Weddell Seas). 
The few records at lower latitudes (<35) 
are at bathyal or abyssal depths. The pre- 
sent data supplement those of Saiz (1995) 
showing this to be a well established inhab- 
itant of the Antarctic Ocean. 


Genus Nephasoma Pergament, 1940 
Nephasoma abyssorum benhami (Stephen, 
1948) 


Material examined.—56°51'S. 34°25'W, 
S17 Oem: His eind= 6 eSepy 19035 SU SNM 
170243; 67°14'S, 70°12'W, 640 m, 3 ind., 
26 Feb 1972, USNM 170420; 73°28/S, 
30°27'W, 3111 m, 4 ind., 13 Mar 1969, 
USNM 170860; 77°32’S. 163°02'W, 624 m, 
Inind:. 95" Feb 1968) USNM, 170963; 
78°24'S, 168°58'W, 565 m, 1 ind., 27 Jan 
1968, USNM 170187 (Fig. 5c). 

Description.—There are 21 specimens 2— 
13 mm long and only 3-6 times longer than 
wide with the smooth white trunks typical 


868 


of this species. The ends of the trunks have 
low skin bodies and may appear rough in a 
few individual, significant papillae are not 
present. The introvent seems to be from 
50—150% of the trunk length and bears dark 
pyramidal hooks 40—100 wm tall that are 
arranged in a irregular spiral manner, not 
random. The pair of retractor muscles orig- 
inate at 5|0—65% of the distance towards the 
posterior end of the trunk, and the longi- 
tudinal muscle layer is fractured into par- 
tially separate bundles in the area around 
the anus in one 10 mm worm. Maturing ova 
are present in one 6 mm specimen that had 
been collected in September. 

The most southerly specimen (USNM 
170187) is quite different looking since it 
seems to have lost the outer layer of cuticle/ 
epidermis such that the underlying papillae 
are now exposed looking like tall, thin, hair- 
like papillae. It is also much larger, the 
trunk being 48 by 7 mm so it is longer and 
thinner that the others. This has clearly ex- 
perienced some post-collection stress. 

Distribution.—The single previous report 
consisted of two worms taken at 66—67° S, 
71 and 138° E, from 540 and 640 m. These 
new records are quite disjunct at 57—77.5° 
S and greatly increase the known range into 
the western longitudes and greater depths. 
One pair of stations is from 3100—3200 m 
at 30.5 and 34.5° W, while the other pair is 
from 600-650 m at 70 and 163° W (Fig. 
Se): 


Nephasoma capilleforme (Murina, 1973) 


Material examined.—74°07'S, 39°38'W, 
650 m, 4 ind., 6 Feb 1968, USNM 170838; 
7718'S, 37°43'W, 1025 m, 1 ind., 6 Mar 
1969, USNM 170844 (Fig. 5c). 

Description.—The five worms are 5—11 
mm long and about 0.5 mm wide so that 
their trunks are 10—20 times longer than 
wide and the introverts are less than half 
the trunk length. The pair of retractor mus- 
cles originates in the middle of the trunk. 
All had been living in tubes of agglutinated 
sand grains and have scattered skin bodies 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


but no papillae on the posterior end. Scat- 
tered small hooks (25—50 jm) are present. 

Distribution.—Widespread in the north- 
ern Atlantic and scattered reports in the 
southern hemisphere, including the Weddell 
Sea at 75° S, and the northern Pacific Ocean 
(to 56° N) at bathyal and abyssal depths 
(900-5000 m). These new records expand 
the southern boundary a small distance to 
Wed Se 


Nephasoma diaphanes (Gerould, 1913) 


Material examined.—There were 589 
specimens from 46 stations (1 station yield- 
ed 472 worms, i.e. 80% of the total) from 
53.2—78.5°S, but with a gap between 66— 
73.5° S. Longitudinally there are scattered 
stations from 27—56° W then a concentra- 
tion at 66—-71° W, and widely scattered sta- 
tions to 180° and a small cluster at 170— 
175° E (Fig. 5c). Vertically the total range 
is from 1—5033 m, but 57% of the stations 
are between 100—1000 m, with 20% <100 
and 23% >1000 m. 

Description.—The vast majority of these 
worms are 1—6 mm long, a few measure 7— 
11 mm. Particular trunk length to width ra- 
tios are characteristic for many species, but 
given the plastic nature of sipunculans this 
value does vary. In 42 measured worms 
71% have lengths 2—5 time the width, 21% 
are 6-8 times longer than wide, but 3 
worms (7%) have lengths 10—13 times the 
width. These last 3 elongate worms ap- 
proach the dimensions of N. capilleforme. 
The skin is generally smooth with low skin 
bodies more concentrated near the ends of 
the trunk. Some worms do have small di- 
gitiform papillae towards the posterior end, 
especially those taken from arenaceous fo- 
raminiferan tests. Introverts are rarely ex- 
tended but most appear to be 50—100% (a 
few up to 150%) of the trunk length. While 
not always present, scattered pyramidal or 
triangular hooks generally 25—40 pm tall, 
can be found. It is very possible that hook- 
less members of this species have been 
identified as small N. eremita. 


VOLUME 114, NUMBER 4 


The single pair of introvert retractor mus- 
cles are fused for about one-half their 
length and they have their origins from the 
body wall 50—75% of the distance towards 
the posterior of the trunk. Developing ova 
were observed in 4 worms 5—6 mm long 
collected in October, February, and April, 
and sperm in a single 5 mm worm collected 
in February. 

Distribution.—Cosmopolitan (from 82° 
N to 50° S) in cold water, most from bathyal 
and abyssal depths (down to 5300 m). This 
material significantly extends its known 
range into and around the Antarctic Ocean. 


Genus Phascolion Théel, 1875 
Subgenus Phascolion (Isomya) Cutler & 
Cutler, 1985 
Phascolion (Isomya) hedraeum Selenka, 
de Man & Biilow, 1883 


Material examined.—The 159 specimens 
are from 23 stations at 54—-61.5° S, plus 1 
at 74.5° S and longitudes 30.5—46° S plus 
1 at 71° W at depths of 70—604 m (Fig. 5d). 

Description.—The trunks are often 
coiled indicating a life within gastropod 
Shells and are 2—25 mm long. The distinct 
holdfast papillae have granular borders that 
may not completely surround the papillae. 
In some worms this hardened protein is eas- 
ily seen but on others may only be seen 
with the compound microscope. The slight- 
ly bent, scattered, spine-like hooks are 55— 
95 wm tall when present. These appear to 
be weakly attached, sometimes just a few 
present, and many worms, often the larger 
ones, have no hooks at all. Around the base 
of the introvert, which is shorter than the 
trunk, are large, densely packed, grey pa- 
pillae, only a very few of these having more 
than a single tip. The two retractor muscles 
originate at the posterior end of the trunk. 
These muscles are usually slightly sub- 
equal in size, the ventral being 75—90% the 
diameter of the dorsal muscle. Developing 
Ova were observed in 7 individuals ranging 
in length from 12—23 mm, having been col- 


869 


lected during the months of February, April 
and June. 

Saiz (1995) identified 5 specimens as P. 
convestitum from the Weddell Sea, now 
considered as P. hedraeum, a closely relat- 
ed species and common representative in 
Antarctic waters. The main difference is the 
shape of the hooks; broad based and re- 
curved in P. convestitum, but blunt and 
spine-like in P. hedraeum. Fig. 4a shows 
variation in this character, which includes 
both shapes. 

Distribution.—Several southern Atlantic 
reports down to 65° S and a few from the 
southern Pacific, near Australia, and also 
off Japan. They live at shelf and slope 
depths, rarely over 800 m in gastropod or 
scaphopod shells. A single specimen from 
4510 m was assigned to this taxon “‘with 
reservations’, and, as stated in that report 
(Cutler 1977), there are several questions 
about the reliability of the station data in 
these Galathea collections. These additions 
add to the one prior Antarctic record (Cutler 
& Cutler 1980) and show a broader, but still 
compact population in this Weddell Sea re- 
gion. 


Subgenus Phascolion (Montuga), Gibbs, 
1985 

Phascolion (Montuga) lutense Selenka, 
1885 


Material examined.—The 709 specimens 
came from 40 stations at latitudes from 53-— 
68° S plus one station at 73.5° S. Longitu- 
dinally, there is a block of stations between 
22—40° W, then scattered records across to 
156° W, plus 3 in the east (108, 158, 164° 
E). This is clearly a deep-water taxon since 
they came from depths of 2160-5790 m, 
50% of the stations and 40% of the speci- 
mens came from depths >4000 m (Fig. 5d). 

Description.—Most of this material was 
10—25 mm long but the total size ranged 
from 4—35 mm. The uniformly smooth skin 
free of visible holdfast papillae, and array 
of small papillae around the base of the in- 
trovert forming a grey cap, characterize this 


870 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
CG eee = 


{ 


Fig. 4. A: Phascolion (Isomya) hedraeum, variation in hook shape. B: Hooks of Phascolion (Montuga) 
lutense. C: Weakly developed holdfast papillae of P. (/.) lutense. Scale bars: A = 50 pm; B = 35 pm; C = 
200 wm. 


VOLUME 114, NUMBER 4 


COO CO Te 5 


tin TEETER 


Fig. 5. 


871 


A-D Distribution maps of the US-collected Antarctic sipunculans. A: (+) Golfingia anderssoni, (0 ) 


G. muricaudata. B: (+) Golfingia margaritacea. C: (+) N. diaphanes; () Nephasoma abyssorum benhami; (@) 
N. capilleforme; (X) Apionsoma murinae murinae; (#) Apionsoma murinae bilobatae. D: (+) Phascolion (Montuga) 
lutense; (©) Phascolion Isomya) hedraeum; (®) Phascolion (Montuga) pacificum; (X) Phascolion sp. 


species. Papillae are present on the trunk 
that might have secreted some peripheral 
protein, but only a trace can be seen in a 
few larger worms (Fig. 4c). It is also the 
only known member of this genus to form 
a stable clay/mud tube as a dwelling place, 
an interesting adaptation to very deep life 
where empty calcareous mollusc shells or 
coral do not exist. The tentacular crown is 
reduced to a membranous ridge that may 


have a few short lobes around the margin, 
another adaptation seen in other deep-sea 
species. Thin spine-like introvert hooks are 
seen only rarely and then in very small 
numbers measuring about 50 pm tall (Fig. 
4b). It seems clear that these are not per- 
manent structures. Living on the anterior 
end of a few worms were small clusters of 
epizoans, probably Entoprocta. 

Internally the retractor muscles are fused 


872 


into a single column for almost their entire 
length, and the single nephridium is often 
quite large. Eggs were observed in only two 
worms 8 and 31 mm long, collected during 
August and October. Sperm were present in 
one 20 mm worm also collected in October. 

Distribution.—A cold-water species 
(1000—6860 m) unknown from lower lati- 
tudes, even in deep water. Widely collected 
in the Southern Hemisphere in all three 
oceans, 36—66° S in the Pacific, 20—32°S 
in the Atlantic, and 32—38° S in the Indian 
Ocean. From northern waters it is recorded 
from the northwestern Pacific and the 
northeastern Atlantic, 47—56° N. This col- 
lection significantly broadens the longitu- 
dinal range at these high southern latitudes. 


Phascolion (Montuga) pacificum Murina, 
QSy7/ 


Material examined.—61°45'S, 61°14'W, 
4758 m, 1 ind, 1 Aug 1962, USNM 
170134; 63°00’S, 82°31'W, 4602 m, 2 ind., 
24 Oct 1963, USNM 170681; 73°28’S, 
30°26’W, 3111 m, 2 ind., 13 Mar 1969, 
USNM 170845 (Fig. 5d). 

Description.—These 5 specimens are 5— 
17 mm long and differ from the other more 
common member of this deep-water sub- 
genus by having holdfast papillae with thin 
but distinct U- or V-shaped hardened bor- 
ders, and they do not construct the clay/ 
mud houses of P. lutense. Both have re- 
tractor muscles almost totally fused into a 
single column, and tentacles reduced to 
lobes. This 17 mm worm was unusually 
thin (2 mm wide) thus not in its natural sau- 
sage shape. Small scattered hooks may be 
present near the tip of the introvert which 
is longer than the trunk in one, but shorter 
in another. 

Distribution.—A bathyal and abyssal 
species (300—6860 m) widespread at high 
latitudes in the northwestern and south- 
western Pacific, the northeastern and south- 
ern Atlantic, and the sub-Antarctic Indian 
Oceans. Two records from lower latitudes 
are the Peru-Chile Trench (5760—6860 m) 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


and 28° N (1760 m) in the eastern Atlantic. 
The only previous Antarctic record was at 
77° E so these new records greatly expand 
the known range to the other side of the 
world in Antarctic waters. 


Phascolion species? 


Material examined.—59°14'S, 69°13'W, 
3738 m, | ind., 6 Oct 1962 USNM 170211 
(Fig. 5d). 

Description.—This single 6 mm worm is 
interesting, but not sufficient for a positive 
identification. It most closely resembles the 
published description of Phascolion usha- 
kovi (Murina, 1974). The similarities are of 
two types; long, thin, dark hooks 200—250 
ym tall and mammiform papillae at both 
ends of the trunk. However, this set of re- 
tractor muscles are not of distinctly differ- 
ent sizes as is the case in P. ushakovi (1.e., 
ventral about 75% the diameter of the dor- 
sal, not the expected 10—20%). This ventral 
muscle has two origins that quickly fuse. 
The mid-trunk region is smooth with no 
holdfast papillae. 

Given that we have only a single small 
specimen, and the lack of congruity in re- 
tractor muscles, we hesitate to assign this a 
species name. Also, the location of our ma- 
terial is far from the only reported location 
of P. ushakovi, i.e., western Australia, at 
330 m. 


Genus Apionsoma Sluiter, 1902 
Apionsoma murinae murinae (Cutler, 


1969) 


Material examined.—59°56'S, 65°18'W, 
3687 m, 1 ind., 10 Aug 1962, USNM 
170204 (Fig. 5c). 

Description.—This single 17 mm speci- 
men is large for this species and has a thin 
introvert about 110 mm long. The posterior 
trunk bears the larger mammiform papillae 
and the tip of the introvert carries rings of 
very small hooks. The nephridia are clearly 
single lobed and while the thin dorsal re- 
tractor muscles are partially hidden under 
the ventral pair they are present. 


VOLUME 114, NUMBER 4 


Distribution.—Widespread in cold water 
(100—5200 m) across the northern Atlantic 
and down to 15° S on the eastern side. In 
the Pacific it has been recorded from the 
Bering Sea and several deep water southern 
locations including the Peru-Chile Trench. 
Recent Indian Ocean records show it to be 
present there at depths from 250—3600 m 
as far south as 38° S. Therefore, this Wed- 
dell Sea specimen is the first for this taxon 
in Antarctic waters. 


Apionsoma murinae bilobatae (Cutler, 


1969) 


Material examined.—54°50'S, 129°48'W, 
LOs5ame 2) ind. 7 Nov 1964, (USNM 
170774 (Fig. 5c). 

Description.—These flask-shaped trunks 
are 8 and 11 mm long with prominent 
mammiform papillae posteriorly. Rings of 
small pale hooks show only very short spi- 
nelets almost as if they had been broken off. 
Since these are larger than most known 
specimens (1—7 mm) it is possible that the 
fragile spinelets wear down with age. This 
is similar to what Popkov (1993) observed 
in his 11 mm specimen from New Zealand 
that he called Apionsoma claviformes. The 
nephridia have the distinctive secondary 
lobe and two pair of introvert retractor mus- 
cles are long and very thin with their ori- 
gins about 75—80% of the distance towards 
the posterior end. 

Distribution.—Known from moderate 
depths in the northern Atlantic Ocean and 
Mediterranean Sea plus scattered reports 
from the eastern and western Indian Ocean, 
and one near New Zealand. Thus, this re- 
cord is the first in the Antarctic region (far 
southern Pacific) for this taxon. 


Discussion 


The taxa discussed above include all that 
have been reported by earlier authors, ex- 
cept these four: 

Golfingia elongata (Hérubel 1906, 
1907a, 1907b) What appears to be three re- 
cords is actually one record repeated three 


873 


times. It is our supposition that this material 
is most likely a few G. margaritacea in 
which Hérubel saw some ordered array of 
hooks. 

Nephasoma confusum (Murina 1957, 
1972) This species is very similar to N. 
abyssorum with moderate sized, dark 
hooks. If the ill-defined spiral order of these 
hooks was not evident these two taxa could 
be easily confused. Hook shape has also 
been used as a diagnostic character. The 
original description of N. confusum by Slui- 
ter (1902) said there is a reinforced rim in 
the hook. However Sluiter illustrated a 
hook with a rim not especially thick. Mu- 
rina (1972) identified material from Antarc- 
tic waters with large hooks and stated that 
their hooks were similar to those drawn by 
Sluiter in 1902. Specimens identified as N. 
confusum sent to one of us (EC) by Murina, 
have large hooks with the suggestion of a 
thin reinforced rim. These hooks are not py- 
ramidal as are the ‘typical’ hook of N. abys- 
sorum. Ditadi & Migotto (1981) stated: 
*“As pointed out by Sluiter (1902) a very 
characteristic feature of this species is the 
chitinous reinforcement of the hooks, a trait 
we have also found and shown in Fig. 
4h4.”” They illustrate several hooks show- 
ing variation, some are only partially rein- 
forced. Cutler & Cutler (1986) and Cutler 
(1994) draw a hook with a thick reinforced 
rim and a little bend in the apex. Saiz & 
Villafranca (1990) found specimens in 
South Spain with the same shape as shown 
in the Cutlers’ work. The typical N. abys- 
sorum hook in Théel (1905) is pyramidal. 
However he drew a specimen with the in- 
trovert out in which the ‘spiral arrange- 
ment’ of the hooks is not obvious. 

In this study we have identified as N. 
abyssorum benhami worms with diverse 
hooks, some having pyramidal edges, oth- 
ers triangular in shape, and others in be- 
tween. It seems that hook shape is more 
variable than previously reported. There- 
fore, we consider it very likely that what 
Murina named N. confusum we would have 
called N. abyssorum benhami. 


874 


Nephasoma eremita (Fischer 1928, Mu- 
rina 1972, Saiz 1995) Given the experience 
of one of us (JISS) it now seems probable 
that these specimens represent a few N. 
diaphanes that lack hooks. 

Phascolion strombus (Fischer 1920, Ste- 
phen 1948, Murina 1972) It is most likely 
that these records are what we are here call- 
ing P. hedraeum. 

Thus, we suggest that this report includes 
all the known members of this phylum liv- 
ing in Antarctic waters. 

Morphology.—The impermanent or de- 
ciduous nature of introvert hooks has been 
observed in a few isolated cases previously, 
but this study affirms the more widespread 
reality of this condition in most of these 
Antarctic species (all three Golfingia spe- 
cies, Nephasoma diaphanes, and all three 
Phascolion species). As a rule hooks are 
less numerous in larger individuals. One 
strong message here is that the absence of 
hooks in a few individuals does not warrant 
the naming of a new species. None of the 
species we encountered are new to science 
and all have been well described from large 
sample sizes, the unnamed Phascolion be- 
ing the single exception. 

Ecology.—Golfingia margaritacea, with 
46% of the specimens, dominates this fau- 
na, but by adding Phascolion lutense with 
22% and Nephasoma diaphanes with 19%, 
we have 87% of the collection in three spe- 
cies. A second tier of species each making 
up 2—5% consists of P. hedraeum, G. an- 
derssoni, and G. muricaudata. The remain- 
ing 5 species if added together comprise 
just 1.1% of the total. 

Of the 456 stations, 395 contained a sin- 
gle sipunculan species, but 58 yielded 2 spe- 
cies and 3 contained 3 species. Eight of the 
11 species were found together with a sec- 
ond. The remaining 3 were the least com- 
mon with only 1—5 representatives each. 

Zoogeography.—The numbers of speci- 
mens examined (3156) and species identified 
(10) in this study are comparable to others 
previously reported from Antarctica (cf. Mu- 
rina 1972, Saiz 1995). All species have been 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


previously collected by other expeditions, 
except for Apionsoma murinae, here record- 
ed for the first time south of the Antarctic 
Convergence. Concerning spatial horizontal 
distribution within Antarctic waters, the 
most ubiquitous species is Golfingia mar- 
garitacea, present along almost the entire 
coastline of Antarctica with a few gaps such 
as off ‘Dronning Maud Land’ (between lon- 
gitude O—45° E) and ‘Marie Byrd Land’ 
(120—150° W). Several other common spe- 
cies (G. anderssoni, G. muricaudata, N. 
diaphanes, P. lutense, P. hedraeum) appear 
a more restricted in those areas heavily sam- 
pled by oceanographic vessels such as the 
Antarctic Peninsula, the Ross and Weddell 
Seas. The remaining species: N. abyssorum 
benhami, N. capilleforme, P. pacificum, and 
A. murinae are represented by only a very 
few specimens from widely scattered loca- 
tions. With regard to the vertical distribution, 
most sipunculan species in the US Antarctic 
collections were found at sub-littoral (1—200 
m) and bathyal (200-3000 m) depths, 
whereas only two, Phascolion lutense and P. 
pacificum, showed a preference for deeper 
water (>2000 m). 


Acknowledgments 


This research was partially supported by 
a grant from the National Museum of Nat- 
ural History, Department of Invertebrate Zo- 
ology, U. S. Antarctic Research Program 
(USARP) funded by the National Science 
Foundation (Contract OPP-9509761). Their 
web site with a searchable database can be 
found at: <http://www.nmnh.si.edu/cgi-bin/ 
wdb/iz/psi/form>. We also acknowledge a 
contribution from the Basque government in 
Spain, contract no. 12736. We are indebted 
to W. Moser and C. Bright (NMNH) for 
their many efforts on our behalf. The staff 
of the Department of Invertebrate Zoology, 
MCZ, Harvard University continue to cheer- 
fully extend essential collaborative support. 


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876 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Appendix.—Station data for sipunculan species Appendix.—Continued. 
found at more than 5 locations. See text for less com- 
mon taxa. Latit. Depth Num. Date 
South Longit. in m. ind. y-m.d. USNM # 
as Depth, }¢ Nuns Date 7736 W04230 0585 69.03.02 170842 


South Longit. in m. ind. y-m.d. USNM # 
SSS eS TVR ENT IND 0746 


Golfingia anderssoni 7750 W04205 0659 
5340 W07015 0307 2 79.04.28 170193 7759 17811 0647 
5357. W03721 0033 75.05.16 170147 7809 W17832 0695 
5403 W03724 0070 75.05.16 170074 7810 W16230 0500 
5414 W03754 0173 75.06.10 170156 7824 WI16858 0565 
5417 W03828 0243 86.12.03 170986 
5422 W03828 0220 86.12.03 170987 
5431 W03535 0265 75.05.19 170149 
5506 W03955 2975 66.02.08 170708 
5629 W02646 2270 75.05.30 170084 
5723 W02634 2325 75.05.26 170151 
5739 W02600 2500 75.05.27 170152 
5855 W15945 0750 64.08.12 170761 
5908 W03649 2817 64.04.10 170172 238 W03802 0137 
6157 W05553 2340 64.03.14 170170 249 W06620 0081 
6203 W06109 1437 62.08.08 170203 2240 W07024 0100 5 
6205 W05823 0048 72.01.15 170399 2342 WO03757 0147 
6207 W05558 1133 63.01.01 170158 242 W07025 0021 
6221 W05848 0082 72.01.16 170976 >>! W03738 0100 3 
6233 W05801 1680 63.01.04 170159 297 W03721 0033 
6241 W05443 0215 64.03.15 170957 400 W03741 0074 
6241 W05443 0215 64.03.15 170253 401 WO03651 0112 
6242 W05610 0430 63.01.02 170220 402 W03740 0058 
6252 W05927 0910 63.01.07 170862 402 W03740 0065 
6256 W06048 0100 72.01.08 170395 5408 W05208 0450 
6326 W06215 0122 69.02.10 170348 411 W03802 0160 
6419 W06259 0045 63.02.06 170820 2412 W03736 0069 
6437 W06251 0655 18 82.03.25 171032 SID WW 037/20 COS OR 
6446 W06404 0028 72.03.10 170438 Stl OS OOE 
6447 W06419 0047 71.12.04 170403 Spl NOB IAN (OON I 
6447 W06407 0055 72.01.03 170392 5412 W03743 0085 
6447 W06410 0208 83.03.20 170125 3414 W03754 0200 
6503 W06354 0365 72.03.01 170421 5422 W03359 0155 
7120 E16955 0439 59.01.25 170803 5424 E15901 0086 
7208 17210 0434 58.01.13 170785 5431 W03535 0265 
7400 W05456 0438 68.02.12 170852 S432 leils202 Cees) 
7432 E16817 0876 68.01.18 170143 5434 W06430 0074 
7457 E17206 0525 68.01.18 170180 5439 W06350 0136 
7458 E16534 #1124 78.02.08 175516 5439 W06420 0048 
7531 W03008 0412 69.02.25 170853 5440 W05705 0126 
7607 W16837 0514 68.02.03 170309 5443 W03649 0188 
7625 W17034 0558 68.01.30 170188 5445 W06410 0022 
7650 W04055 0513 69.03.01 170855 5447 W06404 0040 71.10.29 170379 
7104 16819 0916 68.01.22 170182 5451 W03503 0170 86.12.10 170971 
7105 W03504 0793 69.02.26 170854 5456 W03550 0151 47 75.06.06 170098 
7709 W03858 0851 69.04.05 170843 5500 W06450 0543 2, 7TAMMOSSaOSeH 
7716 W04238 0512 69.03.01 170133 5503 W05857 NR. 13 62.12.04 170216 
7716 W04238 0512 69.03.01 170841 5507 W03547 0131 4 75.06.06 170097 
TQ) Wine  O6ee 67.01.24 170141 5509 W03545 0130 11 75.06.06 170096 
7730 16800 0107 48.01.29 021097 5542 W03539 0240 1 86.12.11 170972 
7731 W16305 0606 68.02.05 170190 5549 W02211 4256 1 66.03.04 170710 
7736 E16609 0249 56.12.26 170780 5606 W06619 0440 2 63.09.18 170769 


78.01.23 175509 
69.03.04 170858 
68.01.25 170185 
78.01.24 175514 
56.01.27 170775 
68.01.27 170138 


Se Se eS LR eS WH 


Golfingia margaritacea 


5306 W06704 0086 
5306 W06704 0086 
5320 W06651 0080 
5331 W07050 0001 
5336 W03735 0310 
5338 W03754 0133 


70.03.05 175492 
70.03.05 175493 
62.09.27. 170210 
69.04.24 170049 
75.05.11 170064 
75.05.13 170068 
75.05.12 170067 
64.02.10 170248 
69.05.13 171090 
75.05.14 170070 
70.04.29 170837 
66.02.07 171004 
75.05.16 170147 
75.05.15 170073 
75.05.17 170075 
75.05.15 170072 
75.05.15 171037 
66.01.30 170274 
86.12.05 170988 
75.06.10 170105 
75.06.10 170104 
75.06.09 170102 
75.06.09 170103 
75.06.09 170101 
75.05.12 170066 
75.05.19 170148 
65.02.10 170764 
75.05.19 170149 
65.02.10 170765 
71.11.07 170382 
71.10.26 170377 
71.10.20 170829 
66.03.14 170276 
75.05.17 170076 
71.05.20 170363 


— 


a NS Be RB OW WO RR RK WNND WR RN RR RK RK WD OR RRP RP RFP NK NWNR KH ON WD Be KP HN Be HB HB HN 


A 
me Ke NO RKB RKP TERE DAA NOR NN OR NKR KK Whe HRY WOK NOD KR KKK NN ND W 


— 


= 


VOLUME 114, NUMBER 4 877 


Appendix.—Continued. Appendix.—Continued. 
Latit. Depth Num. Date Latit. Depth Num. Date 


South Longit. in m. ind. y.m.d. USNM # South Longit. in m. ind. y.m.d. USNM # 


5616 W02730 0290 15 75.06.02 170092 6243 W06102 0073 
5616 W02735 0132 22 75.06.03 170093 6249 W05610 0378 
5619 W02729 0175 27 ©66.03.06 170711 6255 W05927 0915 
5620 W02729 0175 9 66.03.06 171003 6257 WO6051 0154 
5620 W02730 0130 5 75.06.04 170094 6257 W0O6051 0154 
5623 W02723 0185 1 75.06.02 170091 6258 W06041 0087 
5624 W02725 0186 2 75.06.02 170090 6258 W06047 0140 
5642 W02700 0107 75.05.30 170083 6258 W06048 0057 
5643 W02700 0125 75.05.31 170087 6259 W06034 0056 
5644 W02659 0337 75.05.31 170086 6259 W06035 0060 
5644 W02702 0146 75.05.31 170088 6301 W06044 0040 
5645 W02654 0740 75.05.31 170085 6313 W11038 0036 3 
5700 W02610 2742 75.05.22 170150 6314 W05845 0073 
5722 E14937 3299 69.04.19 170320 6314 W0O5845 0073 
5730 W0O5858 3533 62.12.07 170217 6324 W04451 3622 
5739 W02600 2500 75.05.27 170152 6324 W06210 0074 
5741 W02622 0412 75.05.26 170082 6324 W06214 0093 
5839 W06407 0024 71.05.17 170362 6326 W06215 0122 1 
5900 W04900 3867 63.03.08 170677 6351 W06238 0147 
5928 W02703 0093 76.02.27 170061 6351 W06238 0147 
5953. W03219 0600 76.02.25 170869 6351 W06238 0147 
6026 W04622 0105 76.02.16 170108 6400 W06100 0570 
6026 W04623 0155 76.02.16 170109 6402 W06407 0075 
6026 W04628 0110 76.02.16 170110 6402 W06407 0075 
6027 W04550 0103 76.02.15 170106 6406 W11517 4873 
6030 W04643 0126 76.02.16 170058 6409 W06240 0097 
6030 W04643 0129 76.02.17 170059 6413 W06105 0115 
6116 WO5611 0224 62.12.31 170218 6413. W06105 0131 
6117 W04429 0295 76.02.22 170060 6413. W06105 0129 
6119 W0O5609 0230 3 62.12.31 170135 6414 W06158 0260 
6119 W0O5609 0230 62.12.31 170953 6414 W06235 0085 
6125 W05630 0300 64.03.13 170251 6414 W06235 0085 
6202 WO05905 0101 63.01.12 170230 6418 W06258 0170 
6205 W05823 0048 72.01.15 170399 6419 W06259 0045 
6206 WO05600 1200 63.01.01 170219 6419 W06259 0045 


72.01.14 170398 
63.01.02 175502 
63.09.07 170162 
70.03.28 171042 
70.03.28 175490 
70.03.25 170357 
70.03.25 170356 
70.03.25 170355 
82.03.13 171014 
82.04.03 170992 
74.01.12 175517 
61.12.28 170129 
63.01.08 170225 
63.01.08 170226 
63.02.24 170234 
85.03.31 170969 
69.02.10 170449 
69.02.10 170348 
63.01.09 170228 
63.01.09 170229 
63.01.09 175500 
83.03.23 118765 
72.03.01 170432 
72.03.01 170978 
66.04.30 170984 
69.02.10 170448 
71.12.16 170410 
71.12.16 170411 
71.12.16 170412 
82.03.24 171029 
82.03.24 175494 
82.03.24 171031 
82.03.22 171024 
63.02.06 170810 
63.02.06 170819 


N. 
Ne) 


— 


WOreMOmre Be We TOHRK DK OR NABANKP NFP NKR KrF POPPA AN N RR KR WRK NK KON KH AO 
Ne NY RB RN WD BR WB RB YQ BB Mme BB BB BB WB BB OD BB Re Be Ne Re YY 


6218 W05835 0066 71.12.21 170384 6419 W06259 0107 34 =©82.03.22 171023 
6219 WO5911 0044 71.12.19 170414 6419 W06300 0067 5 82.03.22 171022 
6219 WO05911 0044 71.12.19 170415 6420 W06358 0005 4 69.02.03 171038 
6219 WO05911 0044 71.12.19 170866 6421 W06128 0132 1 82.03.23 171025 
6219 WO05911 0044 71.12.19 170867 6421 WO06135 0093 17 =82.03.24 171028 
6219 WO05912 0073 71.12.26 170386 6422 W06128 0105 6 82.03.23 171026 
6221 WO05845 0050 72.01.16 170400 6422 W06128 0106 4 82.03.23 171027 
6221 W05848 0082 72.01.16 175495 6429 W06229 0120 1 82.03.24 171030 
6222 W05912 0670 81.02.21 170128 6434 W06200 0045 5 63.02.09 170821 
6230 W0O05758 1058 63.01.05 170161 6442 W06238 0097 2 71.12.15 170408 
6240 W05445 0265 1 64.03.15 170252 6442 W06238 0098 4 71.12.15 170409 
6240 W05445 0265 64.03.15 175498 6444 W06405 0067 1 72.03.07 170434 
6240 WO05610 0365 1 63.01.02 170222 6445 W06439 0014 1 63.03.01 170825 
6241 W05443 0215 64.03.15 170253 6445 W06408 0193 1 82.03.29 171034 
6241 W05443 0215 64.03.15 170690 6445 W06439 0038 2 63.03.01 170823 
6241 W05443 0215 64.03.15 170999 6445 W06439 0014 1 63.03.01 170824 
6241 W05443 0215 64.03.15 170254 6446 W06404 0021 1 73.02.18 170111 
6241 W05602 0225 83.04.01 170127 6446 W06404 0028 2 72.03.10 170437 
6243 WO06101 0059 72.01.14 170397 6446 W06404 0020 4 73.02.18 170443 


878 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Appendix.—Continued. Appendix.—Continued. 

Latit. Depth Num. Date Latit. Depth Num. Date 

South Longit. in m. ind. y.m.d. USNM # South Longit. in m. ind. y.m.d. USNM # 
6446 W06404 0027 63.01.25 170814 6613 W11038 0038 2 61.12.29 170055 
6446 W06405 0020 71.12.31 170388 6615 E11029 0045 3 58.01.23 170784 
6446 W06406 0224 73.11.08 175513 6617 E11031 0137 7 62.02.12 170052 
6446 W06407 Oll1 72.01.02 170391 6618 E11032 0101 14 61.12.11 170050 
6447 W06407 0027 73.02.20 175508 6621 E11027 0210 33 61.12.06 170053 


6447 W06326 0125 
6447 W06403 0031 
6447 W06404 0037 
6447 W06404 0048 
6447 W06404 0048 
6447 W06405 0017 
6447 W06406 0213 
6447 W06407 0069 
6447 W06407 0100 
6447 W06407 0055 
6447 W06407 0107 
6447 W06407 0108 
6447 W06407 0100 
6447 W06407 0100 
6447 W06407 0077 
6447 W06410 0208 
6447 W06411 0245 
6448 W06330 0056 
6448 W06404 0055 


73.03.11 170122 6621 E11028 0238 
69.01.27. 170057 6622 E11029 £0315 
73.03.06 170118 6622 E11029 0254 
73.03.06 170119 6622 E11029 0247 
73.03.06 171045 6633 E09254 0080 
73.01.27 175510 6637 E16256 0217 
74.10.23 175512 6650 E16432 3515 
73.02.19 170112 6653 E16319 0100 
73.02.20 170114 6718 E11019 0135 1 
72.01.03 170392 6730 W06800 0073 
72.01.26 170402 6730 W06800 0073 
72.02.23, 170417 6730 W0O6800 0074 
72.02.23, 170418 6730 W0O6800 0074 
72.02.23 171043 6848 W06407 0100 
72.03.17 170440 7003 W11451 0366 
83.03.20 170125 7055. E17155 2273 
82.03.29 171035 7122 E17043 0162 
63.01.26 170815 7130 E16930 0250 
72.03.10 170436 7132 E17018 0027 


61.12.03 170056 
61.12.13 170051 
61.12.13 170054 
61.12.13 170870 
58.01.29 170786 
67.02.05 170453 
67.02.05 170451 
74.02.10 170047 
58.01.27 170793 
48.02.22 021091 
48.02.22 021092 
48.02.22 021089 
48.02.22 021093 
56.02.07 170796 
56.02.06 170795 
59.01.18 170801 
68.02.13 170303 
56.02.07 170796 
56.02.06 170795 


WrRereWWRrRONFNN PRP RP RP BPN Pre NP we 


6448 W06407 0100 45 72.01.28 170416 7150 W0O1550 1102 59.01.18 170801 
6448 W06407 0100 S PALA W744} 7206 E17215 0392 58.01.12 170792 
6448 W11430 4886 3 64.01.11 170683 7208 E17210 £0434 58.01.13 170785 
6449 W06205 0075 1 63.03.01 170812 7216 E17018 0134 59.01.12 170799 
6449 W06330 0024 1 70.03.16 170352 7217 E17018 0134 59.01.12 170800 
6449 W06330 0023 1 70.03.16 171041 7304 E17903 0570 68.02.11 170311 
6449 W06333 0053 1 73.03.09 170120 7319 E16915 0100 3 56.02.12 170797 
6449 W06333 0053 1 73.03.09 171046 7359 17051 0597 68.01.13 170296 
6449 W06407 0093 4 82.03.29 171036 7400 W05456 0438 68.02.12 170852 
6452 W06340 0290 1 73.02.19 170113 7401 17853 0257 68.01.14 170298 
6453 W06253 0268 ne oF 00 i ms W708 26) 7406 W03938 0731 68.02.06 170850 
6453 W06333 O115 1 73.03.09 175489 7406 W17911 0256 68.01.05 170300 
6454 W06447 0480 1 72.03.04 170433 7407 W03938 0650 68.02.06 170838 
6504 W06401 0024 3. 72.03.13 170439 7428 W03032 0513 1 69.02.24 170131 
6504 W06456 0355 2 73.03.03 170117 7431 E17010 0320 67.01.16 171008 
6505 W06455 0270 5 73.03.03 170116 7500 E17642 0336 68.01.17 170179 
6512 W06413 0060 1 63.?.? 170817 7531 W0O3008 0412 69.02.25 170853 
6514 W06412 0335 1 82.03.19 171020 7600 E17210 0575 68.01.19 170181 
6514 W06412 0335 62 82.03.19 171021 7600 WO5500 0457 68.02.09 170851 


6514 W06412 0335 1 
6514 W06415 0054 
6514 W06415 0060 
6515 W06412 0295 
6525 E10113 0183 
6554 W06517 0258 
6605 W06521 0086 
6608 W06635 0175 
6610 E11041 0128 
6610 E11641 0036 
6613 E11038 0036 


68.01.20 170305 
68.01.20 170961 
67.01.20 170286 
67.01.20 171009 
72.02.08 170340 
72.02.09 171011 
72.02.09 170341 
67.01.18 170285 
67.01.21 170140 
69.03.01 170855 
68.01.22 170307 


82.03.19 175505 7600 W17648 0567 
82.03.16 171017 7600 W17648 0567 
82.03.15 170980 7602 E17822 0487 
82.03.19 171019 7602 E17822 0487 
48.01.14 170794 7602 E17950 0394 
82.03.16 170981 7602 W17957 0352 
82.03.16 170990 7602 W17957 0352 
82.03.18 171018 7610 E16817 0075 
61.12.29 170872 7630 E17040 0643 
61.12.29 170871 7650 W04055 0513 
61.12.28 170874 7701 E16838 0917 


NNANNRRRRrFPNRYFPNNUON PORK HRP VNR RP RP RFP NKFPNNKP WHR RP WHR RP RP KP NWR KK WW ON 


NNNKF NHK Ke eS OO 


VOLUME 114, NUMBER 4 


Appendix.—Continued. 


Latit. Depth 
South Longit. in m. 
7702 W16644 0411 
7704 E16819 0916 
7706 W15817 0200 
7709 W03858 0851 
7716 W04238 0512 
7119 E16548 0832 
7720. E17239 0663 
7720 W03641 1079 
7727 E16630 0321 
7730 E16800 0102 
7730 E16604 0364 
7730 E16800 0107 
7734 E17443 0728 
7735 E17458 0731 
7736 W04230 0585 
7739 E16616 0315 
7739 W0O4450 0256 
7740 E16614 0382 
7740 W03530 0391 
7740 W03530 0410 
7743 E16621 0419 
7750 E16636 0020 
77150 E16636 0087 
7750 E16637 0020 
7750 E16637 0020 
7750 W04205 0659 
7751 E16634 0124 
7751 E16637 0051 
7751 E16638 0016 
7751 £16638 0021 
77152 E16641 0190 
77152 E16641 0136 
77152 E16641 0129 
77152 ~=£E16641 0128 
77152 £16643 0284 
77152 £16643 0282 
7752. E17733 0770 
7753 E16644 0585 
77153 16649 0585 
7754 W04513 0252 
7758 E17802 0655 
77159 ~E17811 0647 
7820 W17300 0456 
7823 §E17306 0474 
Golfingia muricaudata 

5351 W03738 0100 
5444 W06353 0044 
5506 W03955 2975 
5619 W02729 0175 
6055 W04441 0237 
6125 W04155 0595 
6219 WO05912 0073 
6240 W05610 0365 


Num. 


ind. 


Ar N 


Oo 
Ii) (en) Fs Ne) Way a SS sp TS) [RS TRS) feny I) Me) Ir) It) = 1S) ai 


iS) 
aS 


Ba ee WH PRR Re OR KS WRK NNR ke 


ON Re kK NN Ke 


Date 
y.m.d. 


68.02.04 
68.01.22 
60.12.24 
69.03.05 
69.03.01 
68.01.23 
67.01.24 
69.02.27 
56.02.18 
48.01.29 
56.12.? 
48.01.29 
67.01.25 
68.01.25 
69.03.02 
60.02.19 
58.01.16 
56.11.04 
59.01.28 
59.01.30 
56.12.29 
60.01.03 
58.12.30 
59.12.16 
59.12.31 
69.03.04 
58.02.05 
58.12.30 
61.04.01 
61.05.14 
61.04.14 
61.05.22 
61.10.11 
61.10.31 
61.06.03 
61.11.02 
67.01.25 
60.11.30 
61.03.08 
69.03.03 
68.01.25 
68.01.25 
72.02.18 
68.01.26 


66.02.07 
71.05.11 
66.02.08 
66.03.06 
76.02.22 
64.04.13 
71.12.26 
63.01.02 


USNM # 


170189 
170182 
170811 
170859 
170133 
170183 
170141 
170132 
170798 
021094 
170777 
021097 
170142 
170144 
170856 
170877 
170787 
170778 
170804 
170806 
170782 
170887 
170876 
170885 
170886 


170858 


170788 
170875 
170888 
170895 
170891 
170892 
170893 
170894 
170889 
170890 
170177 
170882 
170884 
170857 
170184 
170185 
170343 
170186 


175501 
170827 
170708 
170711 
175507 
171000 
170386 
170222 


879 
Appendix.—Continued. 

Latit. Depth Num. Date 

South Longit. in m. ind. y.m.d. USNM # 
6241 W0O5751 0892 1 63.01.05 170224 
6241 W05443 0215 1 64.03.15 170999 
6258 W06050 0123 1 70.03.28 170358 
6413 W06105 0118 4 71.12.16 170413 
6442 W06238 0098 2 71.12.15 170409 
6446 W06405 0054 1 72.03.07 170435 
6447 W06407 0143 1 72.01.26 170977 
6447 W06407 0108 1 72.02.23 170417 
6448 W06407 0100 2 72.01.28 170416 
6449 W06251 0075 2 63.02.04 170818 
6452 W06333 0114 1 73.03.09 170121 
6453. W06253 0268 4 73.03.11 170123 
6504 W06456 0355 1 73.03.03 170117 
6635 E09040 0278 2 47.12.30 021096 
7120 E16955 0439 2 59.01.25 170803 
7319 E16915 0100 7 36.0212 170797 
7730 E16800 0107 3 48.01.29 021090 
7736 W04230 0585 4 69.03.02 170856 
77140 W03530 0393 1 59.01.28 170802 
7740 W03530 0410 2 59.01.30 170805 
7740 W03530 0391 1 59.01.28 170804 
7743 E16619 0391 1 60.02.19 170878 
7750 E16636 0020 2 60.01.03 170887 
Nephasoma diaphanes 
5311 W0O7050 0001 1 66.03.27 170277 
5329 E16948 0591 2 68.01.01 170289 
5333 W06457 0121 3 64.02.12 170249 
5335 W06945 0041 1 69.05.17 170124 
5346 W0O7053 0288 472 64.02.06 170982 
5349 W07025 0021 3 70.04.29 170360 
5408 W05208 0450 4 66.01.30 170274 
5427 W0O6612 N.R* 1 63.01.21 170231 
5429 W06350 0840 2 71.10.27 170378 
5434 W06410 0073 1 71.10.20 175488 
5434 W06420 0091 1 71.10.20 170375 
5445 W06100 0001 2 71.05.20 170371 
5445 W06353 0044 11 71.05.11 170361 
5446 W06423 0051 2 71.05.24 170974 
5458 W06503 0247 28 64.02.10 170687 
5459 W10510 4180 1 65.11.20 170704 
5502 W05857 0120 1 62.12.04 170952 
5542 W07035 0174 2 62.08.20 170208 
5606 W06619 0440 1 63.09.18 170680 
5606 W06619 0440 6 63.09.18 170769 
5606 W06619 0436 1 63.09.18 170995 
5624 W02719 0950 2 75.06.02 170089 
5651 W03425 3170 1 63.09.06 170242 
5706 W06321 3975 1 62.07.28 170199 
5800 W07056 4010 1 62.11.03 ?EL311 
5829 E17948 0592 1 68.01.01 170288 
6051 W04255 0300 5 64.04.14 170259 
6056 W05652 2850 1 64.03.13 170689 
6057 W11453 5033 2 64.01.06 170682 


880 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Appendix.—Continued. Appendix.—Continued. 

Latit. Depth Num. Date Latit. Depth Num. Date 

South Longit. in m. ind. y-m.d. USNM # South Longit. in m. ind. y-m.d. USNM # 
6145 WO06114 4728 1 62.08.01 170201 6144 W09952 5046 1 65.07.16 170700 
6442 W06238 0097 2 71.12.15 170830 6145 W06114 4758 1 62.08.01 170200 
6446 W04130 4575 1 69.03.19 170861 6239 W06406 3737 1 62.08.06 170202 
6447 W06410 0198 1 83.03.20 175491 6307 W12812 4682 5 66.11.03 170444 
6535 W06540 0200 1 73.03.01 175511 6353 W10839 4948 46 66.03.27 170983 
6558 W0O6651 0200 1 59.04.04 170809 6406 W11517 4873 2 66.04.30 170984 
7316 W17705 1210 1 68.02.10 170310 6408 W04048 4510 2 64.03.21 170255 
7340 E17517 0519 1 57.11.09 170781 6417 W13013 4645 8 64.06.22 170173 
7406 W03938 0731 2 68.02.06 170130 6417 W13013 4695 2 64.06.22 170757 
7406 W03938 0731 2 68.02.06 170850 6448 W11430 4886 2 64.01.11 170683 
7428 W03032 0513 1 69.02.24 170839 6520 W11730 4845 1 64.06.15 170756 
7604 W17640 0564 1 68.01.21 170306 6537 W12106 4781 1 64.06.17 170262 
7658 W17107 0431 1 68.02.02 170308 6612 W10228 4713 6 64.06.10 170755 
7705 W03504 0743 2 69.02.26 170840 6650 E16432 2515 2 67.02.05 170451 
7705  W03504 0793 9 69.02.26 170854 6748 E15812 Zl) 3 67.02.05 170178 
Vis2 WV SS 0770 1 67.01.25 171002 7328 W03027 3111 6 69.03.13 170860 
7823 W17306 0474 1 68.01.26 1710962 


Phascolion Usomya) hedraeum 
Phascolion (Montagu) lutense 5302 W03740 3080 


— 


63.09.17 170245 


5302 W03740 3080 1 63.09.17 170245 5338 W03802 0137 2 75.05.12 170067 
5403 W03340 2690 21 63.09.09 170160 5342 W03757 0147 3. 75.05.14 170070 
5501 WO9915 3931 1 65.06.05 118738 5344 W03759 0133 3. 75.05.13 170069 
5501 WO9915 2160 4 65.06.05 118728 5346 W07053 0288 2 64.02.06 170982 
5506 W0O3955 2975 2 66.02.08 170708 5351 W03738 0100 78 66.02.07 170707 
5508 W02559 5440 128 63.03.29 170679 5403 W03724 0070 1 75.05.16 170074 
5508 W02559 5440 3. 63.04.29 170758 5406 W03631 0135 1 75.05.19 170078 
5549 W0O2211 4256 9 66.03.04 170710 5411 W03802 0160 1 86.12.05 170988 
5552 W02449 5790 54 63.04.30 170759 5412 W03740 0070 4 75.06.10 170100 
5618 W15613 3694 4 66.11.11 170445 5414 W03754 0200 1 75.05.12 170065 
5653. W03733 3141 1 83.08.28 170240 5422 W03359 0155 40 75.05.19 170148 
5700 W0O2610 2742 39 75.05.22. 170150 5439 WO03851 0267 1 75.06.09 170155 
5723 W02634 2325 109 75.05.26 170151 5451 W03503 0170 1 86.12.10 170971 
5739 WO02600 2500 113 75.05.27 170152 6009 W12709 4375 1 65.10.09 170835 
5806 W12008 4173 1 64.10.31 170772 6022 W04650 0350 3 64.04.15 170697 
5808 W04456 2800 4 63.02.15 170676 6051 W04255 0300 4 64.04.14 170696 
5811 W02548 2400 14 63.05.03 170164 6125 WO4155 0396 3 64.04.13 170258 
5814 W02544 2492 55 63.05.03 170776 6125 W04155 0595 1 64.04.13 170694 
5900 W03945 2825 4 66.02.10 170767 6125 W0O4155 0595 1 64.04.13 170137 
5903. W03649 2817 41 64.04.10 170172 6125 W0O4155 0595 3 64.04.13 170695 
6056 W05652 2850 1 64.03.13 170689 6127 WO4155 0604 1 64.04.12 170257 
6100 WO09502 4941 2 64.10.11 170763 6127 WO4155 0604 1 64.04.12 170692 
6120 W08237 4455 1 63.10.22 170863 6127 W04155 0604 1 64.04.12 170693 
6127 E10800 5043 8 66.04.25 170713 7428 W03032 0513 3 69.02.24 170839 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(4):881—886. 2001. 


Two new species of Nereis (Polychaeta: Nereididae) from the 
Mexican Pacific 


Jesus Angel de Le6n-Gonzalez and Vivianne Solis-Weiss 


(JAL-G) Laboratorio de Zoologia de Invertebrados, Facultad de Ciencias Biol6gicas, Universidad 
Aut6noma de Nuevo Leon, Ap. Postal 5 “F’’, San Nicolas de los Garza, 
Nuevo Le6én, 66451 Mexico; 
(VS-W) Laboratorio de Ecologia Costera, Instituto de Ciencias del Mar y Limnologia, U.N.A.M., 
Ap. Postal 70-305, Mexico, D.E 04510, Mexico 


Abstract.—Two new species of Nereis from the Mexican Pacific are described. 
N. inflata, is characterized by an enlarged dorsal ligule in posterior notopodia 
and long shafted notopodial homogomph falcigers; N casoae belongs to the 
group of species with short dorsal ligules and elongate ventral ligules, but it can 
be differentiated by the presence of long blades in notopodial homogomph fal- 
cigers, with many small teeth on the interior margin and an apical blunt tooth. 


The descriptions included in this study are 
part of an exhaustive revision of specimens 
from the family Nereididae in Mexican 
shores (de Le6n-Gonzalez 1997). So far, 22 
species of the genus Nereis are known from 
the Mexican Pacific. The species newly de- 
scribed herein were collected either by Ma- 
ria Elena Caso in Chivos Island in front of 
Mazatlan harbor, Sinaloa State, Mexico in 
the intertidal zone, by one of us (VSW) in 
soft bottoms of the continental shelf along 
the Sinaloa coast with personnel from the 
Lab de Ecologia Costera, ICML-UNAM, as 
part of the Institutional projects SIPCO and 
CORTES (1981-1982), on board the R/V El 
Puma, of UNAM, and by the first author 
(JALG), those last separated from the en- 
dolithic fauna in sandy rocks collected by 
shrimp trawls in the western coast of the 
Baja California peninsula. 

The type material is deposited in the 
polychaetological collections of both the 
Universidad Autonoma de Nuevo Leon 
(UANL), and the Instituto de Ciencias del 
Mar y Limnologia, U.N.A.M. (CPICML) 


Nereis casoae, new species 
Jena, Nyy 


Material examined.—S55 specimens. Col. 
Dra. Maria Elena Caso, Isla los Chivos, 


Mazatlan, Sinaloa, 9 May 1979 (Holotype 
UANL 4832, 6 Paratypes UANL 4833); 
Project SIPCO I, Stn. A-2(14), 22°17’N, 
106°10'W, 61 m, 23 Apr 1981; Stn. C-1 (1), 
23°37'N, 106°56'W, 40 m, 24 Apr 1981; C- 
2 (7), 23°34'N, 106°57'W, 66 m, 24 Apr 
1981; SIPCO II, Stn. A-2 (2), 22°18'N, 
106°10’W, 66 m, 22 Aug 1981; Stn. C-2 
(2) e235 36)N 5 OT O2UW Zerit 2 3) Aus: 
1981; SIPCO III, Stn. A-2 (2), 22°17'N, 
106°11’W, 74 m, 15 Jan 1982; Stn. C-2 
(17), 23°32’N, 107°08'W, 76 m, 16 Jan 
IOS2- eroject) CORME SE Stnen 33 Cl): 
29°54'N, 114°20'W, 75-80 m, 8 May 1982; 
Stn. 48 (1), 28°13’N, 111°28'W, 54 m, 11 
May 1982. 

Epitokous material examined.—S\IPCO 
II, Stn. A-2 (1). 

Description.—Holotype incomplete with 
71 setigers, 40 mm long, 3 mm wide. Body 
reddish-brown, with dorsal dark band in 
median anterior region, beginning on setig- 
er 2. Prostomium pentagonal, one pair of 
slender frontal antennae. Two pairs of eyes 
in trapezoidal arrangement, anterior pair as 
a lens, posterior oval. Biarticulate palps 
with cylindrical palpostyle. Expanded per- 
istomium, with four pairs of tentacular cirri, 
the longest extending to setiger 4 (Fig. la). 


882 


Pharynx with paragnaths in all areas, ex- 
cept area V, paragnaths arrangement as fol- 
lows: I = 1 small cone, If = 2 cones in a 
triangle; III = 23 cones in a square; [IV = 
22 cones in a cresent; VI = 4 cones in a 
diamond; VII—-VIII = 4 cones in a row. 

Anterior notopodia with dorsal ligules 
triangular, median ligule conical, superior 
lobe reduced as a ridge; neuropodia with 
postsetal lobe rounded, ventral ligule su- 
bulate; dorsal cirrus basally inserted, ventral 
cirrus smaller (Fig. 1b). Median and pos- 
terior parapodia similar to anterior ones, but 
with ventral ligule notoriously elongate. 
Ventral cirrus becoming smaller in posterior 
parapodia (Fig. Ic, d). 

Setae in anterior parapodia arranged as 
follows: notosetae in supracicular position 
homogomph spinigers; supracicular neuro- 
setae homogomph spinigers and _ hetero- 
gomph falcigers strongly denticulate (Fig. 
le); infracicular spinigers and falcigers het- 
erogomph, the latter with a slightly longer 
blade (Fig. 1f). Median and posterior para- 
podia with setae in the following arrange- 
ment: notosetae in supracicular position 
homogomph spinigers and falcigers with 
the anterior region of blade rounded (Fig. 
1g); supracicular neurosetae homogomph 
spinigers and heterogomph falcigers, the 
latter with a short blade, strongly denticu- 
late in inner margin and an apical tooth di- 
rected downwards (Fig. 1h); infracicular 
neurosetae homogomph spinigers and _ het- 
erogomph falcigers, with blade slightly 
larger than supracicular ones (Fig. 11). 

Pygidium absent in this specimen; in an- 
other one, pygidium with terminal anus and 
two longanal cirri. 

Epitokous male.—Specimen complete, 
light yellow, with 67 setigers, 13 mm long 
atokous region 2 mm wide, epitokous re- 
gion 2.5 mm wide. 

Prostomium pentagonal, as long as wide, 
one pair of cirriform antennae, two pairs of 
enlarged eyes, biarticulate palps with 
rounded palpostyles. Peristomium as wide 
as the next two setigers; longest peristomial 
cirrus extends posteriorly to setiger 6. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Body divided in two regions: anterior 
one atokous, first 9 setigers with dorsal and 
ventral cirri slightly modified, all with distal 
cirristyles (Fig. 2a, b). From setigers 11 to 
14, parapodia similar to atokous specimens 
(Fig. 2c). Parapodia from heteronereidid re- 
gion strongly modified, with lamellae as- 
sociated with lobes, ligules and cirri; dorsal 
cirrus in anterior parapodia with up to 9 
ventral crenulations, ventral cirrus short 
(Fig. 2d). Dorsal cirrus smooth in posterior 
parapodia, ventral cirrus almost twice dor- 
sal cirri size (Fig. 2e). Natatory setae from 
modified region with oar-like appendages. 

Discussion.—The presence of the ventral 
elongate ligule is a characteristic seldom 
observed, being only known in Nereis apa- 
lie Wilson (1985) from Australia, N. fau- 
chaldi de Le6n-Gonzalez & Diaz-Castafie- 
da (1998), from western Mexico and prob- 
ably N. latrescens Chamberlin (1918) from 
California, since in the latter the ventral lig- 
ule is not so elongate as in the former spe- 
cies but more so than in other species of 
Nereis. This structure is present in N. ca- 
soae, new species, as in the species for- 
merly mentioned, but it can be differenti- 
ated by the presence of the long blade in 
notopodial homogomph falcigers, with 
many small teeth on the interior margin, 
and an apical blunt tooth; in N. apalie and 
N. latrescens, for the above mentioned fal- 
ciger, the blade is short and with 5 and 3 
teeth, respectively, whereas in N. fauchaldi 
the blade is short and smooth. 

Etymology.—This species is dedicated to 
Dr. Maria Elena Caso who collected part of 
the material which was used in the descrip- 
tion. 

Distribution.—This species is known 
from the Mazatlan area (Sinaloa State), and 
two localities in the Gulf of California 

Habitat.—Among algae fixed to coral- 
line rocks, and in sandy bottom of the Gulf 
of California. 


Nereis inflata, new species 
Fig. 3 
Material examined.—7 specimens. Col. 
J. A. de Le6n-Gonzélez, in front of Punta 


VOLUME 114, NUMBER 4 883 


Fig. 1. Nereis casoae, new species: a. Anterior region, dorsal view; b. Parapodium, setiger 10, anterior view; 
c. Parapodium, setiger 37, anterior view; d. Parapodium, setiger 71, anterior view; e. Notopodial heterogomph 
supracicular falciger from setiger 10; f. Neuropodial infracicular heterogomph falciger from setiger 10; g. No- 
topodial homogomph falciger from setiger 71; h. Neuropodial supracicular heterogomph falciger from setiger 
37; 1. Neuropodial infracicular heterogomph falciger from setiger 37. Scale bars: a = 1 mm; b—c = 250 wm; d 
= 150 pm; e-1 = 30 pm. 


San Juanico, Baja California Sur, 26°13’N; 56 setigers, 11 mm long, 2 mm wide, in- 
112°32'W, 30 m, 28 Feb 1989 (Holotype cluding setae; body reddish, prostomium 
UANL 4834, 5 Paratypes UANL 4835, 1 strongly pigmented, posterior region of an- 
Paratype CPICML). terior setigers with transverse dark narrow 

Description.—Holotype complete, with band. Prostomium pentagonal, a pair of 


884 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. 


Nereis casoae, new species (epitoke): a. Parapodium, setiger 1, anterior view; b. Parapodium, setiger 


5, anterior view; c. Parapodium, setiger 10, anterior view; d. Parapodium, setiger 20, anterior view; e. Parapo- 
dium, setiger 50, anterior view. Scale bars: a = 100 wm; b-e = 150 pm. 


slender antennae which do not extend to an- 
terior margin of palps. Two pairs of round- 
ed small eyes. Biarticulate palps, conical 
palpostyle. Prostomium as large as the next 
setiger; 4 pairs of tentacular cirri, the lon- 
gest extending posteriorly to setiger 4 (Fig. 
3a). 

Pharynx with paragnaths in following ar- 
rangement: I = 2 cones in a row; II = 11 
cones in 2 rows; III = 11 cones in an oval 


group; IV = 17 cones in a crescent; V = 6 
cones in a group; VI = 6 in a stellate group; 
VII-VIII = 60 cones in 3 rows. 

Anterior parapodia with notopodia 
formed by one median and one dorsal con- 
ical ligules, and subulate superior lobes; 
neuropodia formed by a postsetal mammil- 
liform lobe and a digitiform ventral ligule. 
Dorsal cirrus slender inserted in the median 
posterior region, ventral one inserted basal- 


VOLUME 114, NUMBER 4 


885 


Fig. 3. 


Nereis inflata, new species: a. Anterior region, dorsal view; b. Parapodium, setiger 10, anterior view; 


c. Parapodium, setiger 23, anterior view; d. Parapodium, setiger 41, anterior view; e. Neuropodial infracicular 
heterogomph falciger from setiger 23; f. Homogomph notopodial falciger from setiger 41. Scale bars: a = 1 


mm; b—d = 150 wm; e-f = 15 pm. 


ly, smaller (Fig. 3b). Median parapodia 
with reduced superior lobes, dorsal cirrus 
inserted medially (Fig. 3c). Posterior noto- 
podia with expanded dorsal ligule, superior 
lobe absent; neuropodia with reduced post- 
setal lobe. Dorsal cirrus inserted distally 
(Fig. 3d). 

Setae in anterior parapodia in the follow- 
ing arrangement: notosetae supracicular 


homogomph spinigers; neurosetae supraci- 
cular homogomph spinigers and _ hetero- 
gomph falcigers; infracicular setae hetero- 
gomph spinigers and falcigers, the latter 
with apical tooth directed downwards (Fig. 
3e). Median and posterior parapodia with 
setation similar to those of anterior para- 
podia; they only differ in the notopodium 
where in addition to the homogomph spi- 


886 


nigers homogomph falcigers are present, 
with slender blades strongly denticulate in 
interior margin, with a small distal tooth di- 
rected downwards (Fig. 3f). 

Pygidium with anus terminal and with 
two long ventral cirri. 

Discussion.—Nereis inflata, new species, 
belongs to the group of species with noto- 
podial superior lobes expanded in posterior 
setigers and notopodial homogomph falci- 
gers with long and strongly denticulate 
blades; this group includes: N. callaona 
(Grube, 1857) from Peru, N. lamellosa Eh- 
lers, 1868 from the Adriatic sea and N. pro- 
fundi Kirkegaard, 1956 from western Afri- 
ca; however, only in N. lamellosa is the su- 
perior lobe present in notopodia of anterior 
setigers, but in contrast to N. lamellosa, in 
N. inflata an apical tooth directed down- 
wards is present in notopodial homogomph 
falcigers. On the other hand, in N. inflata 
in area V of the pharynx there are 6 cones 
in a group, whereas in N. lamellosa there 
are 0 to 3 cones in a triangle. According to 
Ehlers (1868), in this species 10 small 
cones are present in an itregular arrange- 
ment on area VI, whereas in the N. inflata 
6 cones are present in a perfect stellate 
shape. 

Etymology.—The specific name “‘infla- 
ta” derives from the Latin “inflatus’” (= 
puffed or swollen) in reference to the shape 
of the notopodial superior lobes in posterior 
parapodia. 

Distribution.—Endemic. Only known 
from one locality in the western coast of 
Baja California Sur. 

Habitat.—The species was collected in 
the cavities of sandy rocks obtained in a 
shrimp trawl. 


Acknowledgments 


We would like to thank M.Sc. Francisco 
Solis Marin for allowing us to examine and 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


use the material collected by Dra. Maria 
Elena Caso, and deposited in the Labora- 
torio de Equinodermos del ICMyL UNAM, 
as well as the personnel of CIBNOR who 
participated in the oceanographic cruise 
MARSEP XVI to the western coasts of 
Baja California. Dr. M. Hendricks, head of 
the projects SIPCO and CORTES is also 
thanked for having invited one of us (VSW) 
to participate in these projects. Part of this 
study was funded by the “‘Comisi6n Na- 
cional para el conocimiento y uso de la Bio- 
diversidad’”” (CONABIO, Project H-011). 


Literature Cited 


Chamberlin, R. V. 1918. Polychaetes from Monterey 
Bay.—Proceedings of the Biological Society of 
Washington 31:399—407. 

de Leon-Gonzalez, J. A. 1997. Nereidos (Polychaeta: 
Nereidae) de los litorales mexicanos: Sistema- 
tica, Biogeografia y Alimentacion. Unpublished 
Tesis Doctoral, Universidad Autonoma de Nue- 
vo Leon, 298 pp. 

, & V. Diaz-Castaneda. 1998. Two new species 
of Nereis (Polychaeta: Nereididae) from Todos 
santos Bay, Ensenada, Baja California, Mexi- 
co.—Proceedings of the Biological Society of 
Washington 111:823-828. 

Ehlers, E. 1868. Die Borstenwiiemer (Annelida: Chae- 
topoda) nach Sy stematischen und Anatomisch- 
en Untersuchungen Dargestellt. Leipzig. Wil- 
helm Engelmann. xxiv + 748 pp. 24 pls (pp. 
1—268, pls. 1-11 published in 1864). 

Grube, A. E. 1857. Annulata Oerstediana. Enumeratio 
Annulatorum, quae in itinere per Indiam occi- 
dentalem et Americam centralem annis 1845— 
1848 suscepto legit cl. A.S. Oersted, adjectis 
speciebus nonnullis a cl. H. Kroyer er in itinere 
ad Americam meridionalem collectis. Pt. 2. Vi- 
densk.—Medd.dansk Naturh Foren. 2:158—166. 

Kirkegaard, J. B. 1956. Benthic Polychaeta from 
depths exceeding 6000 m.—Galathea Reports 
2:63—78. 

Wilson, R. S. 1985. Nereis and Platy nereis (Poly- 
chaeta: Nereididae) from Victoria with de- 
scription of a new species of Nereis.—Pro- 
ceedings of the Royal Society of Victoria 97: 
129-138. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(4):887—896. 2001. 


The genus Pseudoxomysis (Crustacea: Mysidacea: Mysidae: 
Leptomysini), with description of a new species from the Timor Sea 


Masaaki Murano 


Institute of Environmental Ecology, METOCEAN Environment Inc., Riemon 1334-5, 
Ooigawa-cho, Shida-gun, Shizuoka 421-0212, Japan 


Abstract.—Two species of Pseudoxomysis are discussed. The present occur- 
rence of P. caudaensis in coastal waters of Indonesia is only the second record 
for the species since Nouvel established it based on specimens from Vietnam. 
A new species, Pseudoxomysis incisa, is described from the Sahul Shelf, Timor 
Sea. The new species is distinguished from the other species of the genus, P. 
caudaensis, by the smooth body, the third segment of the mandibular palp 
furnished with two series of setae on the outer margin, the endopod of the 
fourth male pleopod extending to the distal end of the fourth exopod segment, 
and the endopod of the uropod armed with 50 spines on the inner margin. 


The genus Pseudoxomysis was estab- 
lished by Nouvel (1973) to accommodate 
P. caudaensis collected from near Nha 
Trang, Vietnam. At that time he moved 
Doxomysis longiura Pillai, 1963, to this ge- 
nus. Since then, no further reports of this 
genus have been presented. The Pseudox- 
omysis specimens in my possession are 
composed of two species, P. caudaensis 
Nouvel, 1973, and an undescribed species. 
Those identified as P. caudaensis were col- 
lected in Tegal, located in the northern coast 
of Java Island, Indonesia, by Mulyadi and 
were presented in my study. The specimens 
of the supposed new species were collected 
off the Sahul Shelf in the Timor Sea during 
a cruise (KH-72-1) of Hakuho Maru of the 
Ocean Research Institute, University of To- 
kyo. This latter species is described herein. 
All the specimens examined are deposited 
in the National Science Museum, Tokyo 
(NSMT). 


Systematics 
Genus Pseudoxomysis Nouvel, 1973 
Pseudoxomysis Nouvel, 1973: 131-132. 


Diagnosis.—Carapace produced anteri- 


orly into triangular rostral plate with round- 
ed apex. Antennal scale lanceolate with 
rounded apex, setose all around. Labrum 
provided with strong frontal process. Distal 
segment of endopod of maxilla expanded 
distally, shorter than broad, distal margin 
nearly straight, with about 10 similar-sized 
strong spines, outer margin naked, clearly 
longer than inner. Endopods of third to 
eighth thoracic limbs with carpopropodus 
divided into 3 subsegments by transverse 
articulation, terminal claw setiform. Exo- 
pod of fourth pleopod of male 7-segmented; 
ultimate segment small, with 2 simple setae 
on terminal end, penultimate and antepen- 
ultimate segments armed with strong mod- 
ified seta. Endopod of fourth pleopod of 
male 6-segmented, considerably shorter 
than exopod, without modified setae. Fe- 
male pleopods rudimentary. Female with 3 
pairs of oostegites. Endopod of uropod 
armed with spines along inner margin. Tel- 
son with deep apical incision furnished with 
spinules on either side and pair of plumose 
setae arising from anterior end. Lateral mar- 
gin of telson with spines throughout. 

Type species.—Pseudoxomysis caudaen- 
sis Nouvel, 1973. 

Remarks.—Nouvel 


(1973) mentioned 


888 


three major characters that distinguish 
Pseudoxomysis from the closely allied ge- 
nus, Doxomysis, and its related genera, as 
follows; the labrum with a strong frontal 
process, the distal endopod segment of the 
maxilla with the outer margin longer than 
the inner, and the endopods of the third to 
eighth thoracic limbs with the carpopropo- 
dus divided into 3 subsegments by the 
transverse articulation. 

When Nouvel (1973) established Pseu- 
doxomysis, he suggested that Doxomysis 
longiura Pillai, 1963, should be transferred 
to this genus, because D. longiura, de- 
scribed and illustrated by Pillai (1963, 
1964), was in agreement with this genus in 
the characteristics of the maxilla and tho- 
racic endopods, and with respect to the la- 
brum he received a private letter from Pillai 
that the lip is anteriorly prolonged into a 
spine though short and apically broad and 
rounded (Nouvel 1973). In 1973, however, 
Pillai described and illustrated this species 
in detail, but omitted to describe the la- 
brum. Liu & Wang (1986) also examined 
D. longiura specimens from northern areas 
of the South China Sea and stated that the 
anterior margin of the labrum was not 
armed with a spine. 

At least, the labrum in D. longiura lacks 
such a strong frontal process as seen in P. 
caudaensis. Furthermore, D. longiura is 
different from Pseudoxomysis in having a 
notable papilla on the eyestalk and fine spi- 
nules between proximal two lateral spines 
of the telson. Considering these circum- 
stances, it is judged that the transfer of D. 
longiura to Pseudoxomysis should be can- 
celed. As a result, the genus Pseudoxomysis 
comprises only two species which are de- 
scribed in this paper. 


Pseudoxomysis caudaensis Nouvel, 1973 
Biesale2 


Pseudoxomysis caudaensis Nouvel, 1973: 
132-141, figs. 1-25. 


Material.—(NSMT-Cr 13404) 1 adult fe- 
male with embryos (5.4 mm), | adult male 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(4.5 mm), 1 immature female with half- 
grown marsupium (4.4 mm) and 1 juvenile 
(2.7 mm); Tegal, northern coast of central 
Java, Indonesia, 3 June 1994, plankton net, 
collected by Mulyadi. 

Description.—Carapace produced ante- 
riorly into triangular rostral plate with 
rounded apex extending to proximal third 
of first segment of antennular peduncle 
(Fig. 1A, B), posteriorly shortened, last 2 
thoracic somites visible dorsally. Anterolat- 
eral corner of carapace rounded. 

Eye slightly compressed dorso-ventrally, 
1.5 times as long as broad; cornea devel- 
oped, as broad as eyestalk; eyestalk 1.5 
times longer than cornea, without papilli- 
form process on dorsal surface (Fig. 1A, B). 

Antennular peduncle of female more 
slender than that of male; first segment 
twice as long as broad, armed with several 
setae at outer distal corner and | seta at 
inner distal corner; second segment short, 
wider than long, armed with 1 long seta 
near distal end of inner margin and | short 
seta on middle of outer margin; third seg- 
ment connected with second obliquely, as 
long as first, twice as long as broad, armed 
with 1 long seta on middle of inner margin 
and | short and 2 long setae at inner distal 
corner (Fig. 1A). In male, first peduncular 
segment 1.5 times as long as broad, armed 
with several setae at outer distal corner and 
1 seta at inner distal corner; second segment 
short, armed with | short seta on middle of 
outer margin, no setae on inner margin; 
third segment longer than first, 1.5 times as 
long as broad, armed with 1 long and 1 mi- 
nute setae at inner distal corner (Fig. 1B). 

Antennal scale extending slightly beyond 
distal margin ‘of antennular peduncle in fe- 
male and to distal end in male, lanceolate 
with rounded apex, more than 6 times as 
long as widest part at proximal fourth; outer 
margin nearly straight, apical suture at 
about distal tenth (Fig. 1C). Antennal pe- 
duncle barely reaching middle of scale, 3- 
segmented, second segment longest (Fig. 
1C). Antennal sympod with | stout spine at 
anterolateral corner (Fig. 1C). 


VOLUME 114, NUMBER 4 


Mandible with masticatory edge devel- 
oped; palp narrow, third segment half as 
long as second, with 7 feathered setae on 
distal third of outer margin (Fig. 1D). Max- 
illule with outer lobe armed with 12 stout 
spines on distal margin and 3 setae on ven- 
tral surface; inner lobe armed with 3 strong 
and 1 slender setae on distal margin, 4 setae 
on outer margin, 2 setae on inner margin 
and 4 setae on ventral surface (Fig. 1E). 
Maxilla with second segment of endopod 
greatly expanded distally, shorter than 
broad; outer margin naked, 1.5 times longer 
than inner, so that distal margin clearly in- 
clined towards mouth; distal margin very 
slightly concave, nearly 3 times as broad as 
at base, armed with 9 strong spines and 2 
setae; inner distal corner with 6 barbed se- 
tae (Fig. 1F). Labrum with strong frontal 
process (Fig. 1G). 

Endopod of first thoracic limb robust; 
lobe from basis rather small, armed with 
many stout plumose setae; preischium and 
ischium expanded medially, wider than 
long; merus relatively slender, twice as long 
as broad; dactylus wider than long, fur- 
nished with stout setae (Fig. 1H). Endopod 
of second thoracic limb rather slender; mer- 
us as long as carpopropodus and dactylus 
combined; dactylus as long as broad, armed 
with setae arranged radially (Fig. 11). En- 
dopods of third to eighth thoracic limbs lost 
in all specimens. Exopods of first to eighth 
thoracic limbs with flagelliform portion 8- 
or 9-segmented; basal plate small, with out- 
er distal corner pointed (Fig. 1H, I. 

First to fifth abdominal somites subequal, 
sixth somite 1.5 times longer than fifth. 
Second to sixth abdominal somites hispid. 
Each of second to fifth abdominal somites 
with small process along ventral median 
line. 

Pleopods of male biramous, natatory. 
First pleopod with endopod reduced into 
short, unsegmented lobe, exopod 7-seg- 
mented (Fig. 2A). Second and third pleo- 
pods with 7-segmented exopod and 6-seg- 
mented endopod, exopod longer than en- 
dopod. Exopod of fourth pleopod slightly 


889 


less than twice as long as endopod, 7-seg- 
mented, ultimate segment small, with pair 
of simple setae on terminal end; penulti- 
mate segment 3 times as long as ultimate 
segment, with | strong seta overreaching tip 
of terminal setae for half of its length; an- 
tepenultimate segment shorter than suc- 
ceeding one, with | strong seta extending 
slightly beyond apex of terminal setae; each 
of proximal 4 segments with pair of ordi- 
nary plumose setae (Fig. 2B). Endopod of 
fourth pleopod 6-segmented, only extend- 
ing to distal margin of third segment of ex- 
opod, without modified setae (Fig. 2B). En- 
dopod of fifth pleopod 6-segmented, first 
segment with triangular lobe tipped with 
seta on outer margin in addition to usual 
side lobe; exopod 7-segmented, longer than 
endopod (Fig. 2C). 

Endopod of uropod overreaching distal 
end of telson for %4 of its length, armed 
along inner margin with about 37 spines of 
variable length (Fig. 2D, E). Exopod of uro- 
pod overreaching apex of telson for more 
than % of its length (Fig. 2E). 

Telson (Fig. 2E, F) as long as last ab- 
dominal somite, 1.7 times as long as broad 
at base, abruptly narrowing towards proxi- 
mal third, parallel-sided in middle third and 
then gradually narrowing distally. Lateral 
margin armed with about 21 spines 
throughout, spines longer in female than in 
male; each apex with single stout spine. 
Apical cleft % as long as telson and U- 
shaped in female, 4% as long and V-shaped 
in male, furnished on either side with 2 dis- 
tal spines and 16—18 spinules; pair of plu- 
mose setae arising from anterior end of 
cleft. 

Remarks.—The present specimens agree 
in main characteristics with the description 
of Pseudoxomysis caudaensis Nouvel, 
1973, but some minor differences are ap- 
parent. The endopod of the fourth male ple- 
opod is slightly less than half as long as 
exopod and extends to the distal end of the 
third segment of the exopod in the present 
specimen, while in the type specimens it is 
slightly more than half as long as exopod 


890 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Pseudoxomysis caudaensis Nouvel, A, C—I, adult female; B, adult male. A, anterior end of adult 


female; B, anterior end of adult male; C, antenna; D, mandible and mandibular palp; E, maxillule; EK maxilla; 
G, labrum; H, first thoracic limb; I, endopod of second thoracic limb. 


VOLUME 114, NUMBER 4 


= 


SSS 


f L 
Ya 

\ 
M NX 
a 
h \ 
\ 
4 
tp Alp, \ 
, 
4 NG \N 
Na 
WAN 
ya 


0.5 mm 1A—F 


Fig. 2. Pseudoxomysis caudaensis Nouvel, A—C, FE adult male; D, E, adult female. A, first pleopod of male; 
B, fourth pleopod of male; C, fifth pleopod of male; D, endopod of uropod; E, uropod and telson, female; EF 


telson, male. 


892 


and extends to the middle of the third seg- 
ment. In the type specimens the telson is 
narrower and several proximal lateral 
spines arise from the dorsal surface rather 
than from the lateral margin, while in the 
present specimens the spines arise from the 
lateral margin. 

Distribution.—The type specimens were 
collected from Cauda Bay near Nha Trang, 
Vietnam, and the present specimens from 
Tegal, northern coast of the central Java, In- 
donesia. This species seems to be a coastal 
form in Southeast Asia. 


Pseudoxomysis incisa, new species 
Figs. 3, 4 


Type  series.—Holotype (NSMT-Cr 
13401), adult male (6.4 mm); allotype 
(NSMT-Cr 13402), adult female with em- 
bryos (5.4 mm); paratypes (NSMT-Cr 
13403), 6 adult males (5.2—5.8 mm), 6 adult 
females (4.7—5.9 mm), 3 of which (4.7, 5.6, 
5.9 mm) bear embryos; Sahul Shelf, 
ZW S53 IWS By ite) WATS 
129°41.8'E; 49-52 m; 24 June 1972; col- 
lected with plankton net installed in mouth 
of 3-m beam trawl during Hakuho Maru 
Cruise (KH-72-1). 

Other material.—17 adult males (5.2—6.4 
mm), 13 adult females (4.7—5.9 mm), 4 im- 
mature males, 9 immature females; collec- 
tion data same as type series. 

Description.—Body smooth. Carapace 
produced frontally into triangular rostral 
plate with narrowly rounded apex extend- 
ing to proximal third to middle of first seg- 
ment of antennular peduncle, lateral margin 
of rostrum slightly concave or straight (Fig. 
3A, B). Anterolateral corner of carapace 
rounded. Posterior margin slightly emargin- 
ate, leaving last 2 thoracic somites exposed 
dorsally. 

Eye relatively large; cornea occupying 
about half of whole eye, slightly wider than 
eyestalk; eyestalk hispid on proximal cor- 
ners, without papilliform process on dorsal 
surface (Fig. 3A, B). 

Antennular peduncle of male more robust 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


than that of female. In male first segment 
with several setae on anterolateral corner 
and | seta on anteromedial corner, second 
segment shortest, third segment about as 
long as first, 1.3 times longer than broad, 
with 3 setae on anteromedial corner (Fig. 
3A). In female first segment more than 
twice as long as broad, second segment 
with 1 seta on anteromedial corner, third 
segment slightly shorter than first, slightly 
less than twice as long as broad, with 1 seta 
on middle of medial margin and 6 setae on 
anteromedial margin (Fig. 3B). 

Antennal scale lanceolate with rounded 
apex, extending beyond distal end of anten- 
nular peduncle for 1/7 of its length in male 
and for 1/6 in female, about 5.5 times as 
long as maximum breadth at about proxi- 
mal third, with suture at distal tenth, setose 
all around; inner margin convex; outer mar- 
gin straight (Fig. 3A—C). Antennal peduncle 
short, reaching middle of antennal scale, 
second segment longest. Antennal sympod 
with spine at anterolateral corner. 

Mandible with masticatory edge devel- 
oped. Mandibular palp narrow, third seg- 
ment half as long as second, outer margin 
armed with 2 series of setae, 10 proximal 
setae becoming longer distally, feathered on 
proximal half, 5 distal setae of equal length 
feathered along whole length (Fig. 3D). 
Maxillule, outer lobe armed with 11 stout 
spines on distal margin and 3 setae on ven- 
tral surface; inner lobe armed with 3 strong 
and | slender setae on distal margin, 4 setae 
on outer margin, 3 setae on inner margin 
and 7 setae on ventral surface (Fig. 3E). 
Maxilla with second segment of endopod 
greatly expanded distally, outer margin lon- 
ger than inner, naked, distal margin trun- 
cate, 3 times broader than basal margin, 
armed with 9 stout, naked spines, outer 2 
spines, especially outermost one, consider- 
ably longer than others (Fig. 3F). Labrum 
armed on frontal margin with pointed, long, 
robust process (Fig. 3G). 

Endopod of first thoracic limb robust; 
lobe from basis armed with many stout plu- 
mose setae, preischium and ischium ex- 


VOLUME 114, NUMBER 4 893 


Fig. 3. Pseudoxomysis incisa, new species, A, C—I: holotype; B: allotype. A, anterior end of adult male; B, 
anterior end of adult female; C, antennal scale; D, mandible and mandibular palp; E, maxillule; EK maxilla; G, 
labrum; H, first thoracic limb; I, endopod of second thoracic limb. 


894 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


2 —S 
ae 


Fig. 4. Pseudoxomysis incisa, new species, A, allotype; B, one of male paratypes; C—G, holotype. A, endopod 
of third thoracic limb without dactylus; B, endopod of seventh thoracic limb; C, fourth pleopod of male; D, 
distal part of exopod of fourth male pleopod; E, fifth pleopod of male; K endopod of uropod; G, uropod and 
telson. 


VOLUME 114, NUMBER 4 


895 


Table 1.—Morphological differences between Pseudoxomysis caudaensis Nouvel, 1973 and P. incisa, new 


species. 


P. caudaensis 
Nouvel, 1973 


Abdomen 
Mandibular palp 


Hispid 


Fourth pleopod of male 


segment 
Uropod 


of its length 


panded medially, wider than long, merus 
relatively slender, twice as long as broad, 
dactylus wider than long, hirsute (Fig. 3H). 
Endopod of second thoracic limb rather 
slender, merus as long as carpopropodus 
and dactylus together, dactylus longer than 
broad, with terminal claw strong (Fig. 31). 
Endopods of third to eighth thoracic limbs 
slender, with carpopropodus divided into 3 
subsegments by transverse articulations 
(Fig. 4A, B). 

First 5 abdominal somites subequal, sixth 
somite 1.3 times longer than preceding one. 

Male pleopods developed, biramous. 
First pleopod with unsegmented endopod 
and 7-segmented exopod. Second and third 
pleopods with 7-segmented exopod and 6- 
segmented endopod, endopod shorter than 
exopod. Exopod of fourth pleopod 1.5 
times longer than endopod, 7-segmented; 
antepenultimate segment shorter than pre- 
ceding one, armed with 2 setae, one short, 
arising from outer distal corner, the other 
markedly robust, arising from near distal 
end and reaching tip of terminal setae, 
curved inwardly; penultimate segment 1.5 
times longer but narrower than preceding 
one, armed with 2 unequal setae on distal 
margin, shorter one as long as ultimate seg- 
ment, longer one 1.5 times longer than seg- 
ment supporting it, straight, with short 
spine-like setae on distal half; ultimate seg- 


Third segment with a single series of 
setae on outer distal margin 

Exopod slightly less than twice as long 
as endopod; endopod extending to 
middle to distal end of third exopod 


Endopod armed with 37 spines along 
inner margin; exopod overreaching 
distal end of telson for more than % 


P. incisa, 
new species 


Smooth 

Third segment with 2 series of setae on 
outer margin 

Exopod 1.5 times as long as endopod; 
endopod extending to distal end of 
fourth exopod segment 


Endopod armed with about 50 spines 
along inner margin; exopod over- 
reaching distal end of telson for near- 
ly half of its length 


ment small, nearly % of preceding one in 
length, armed with 2 naked, equal-sized se- 
tae on distal end (Fig. 4C, D). Endopod of 
fourth pleopod extending to distal margin 
of fourth segment of exopod, 6-segmented, 
without modified setae, first 4 segments 
with naked accessory seta (Fig. 4C). Fifth 
pleopod with 7-segmented exopod and 6- 
segmented endopod; first segment of en- 
dopod with slender lobe tipped with seta in 
addition to usual side lobe; third and fourth 
segments of endopod with naked accessory 
seta (Fig. 4E). Side lobe on second to fifth 
endopods broad (Fig. 4C, E). 

Uropod slender, setose all around; endo- 
pod extending beyond distal end of telson 
for % of its length, furnished with about 50 
spines on inner margin from statocyst re- 
gion to apex; exopod overreaching telson 
for nearly half of its length (Fig. 4E G). 

Telson slightly shorter than last abdomi- 
nal somite, 1.7 times as long as maximum 
basal width, abruptly narrowing near base, 
then gradually towards apex. Lateral mar- 
gin of telson armed with 16—19 acute spines 
arranged sparsely in proximal half and 
densely in distal half, proximalmost spine 
stout, arising at widest part, proximal sec- 
ond and third spines arising from dorsal 
surface near margin. Each apical lobe 
armed with 1, occasionally 2, strong spines 
on apex and 2 spines near distal end of cleft 


896 


margin. Cleft extending to about % of telson 
length, armed with 16 or 17 spinules on e1- 
ther side, anterior end with pair of plumose 
setae between which small sinus is present 
(Fig. 4G). 

Etymology.—The specific name incisa is 
from the Latin incisus = cut into, derived 
from caedo = to cut, in reference to the 
apical cleft of the telson. 

Remarks.—The present new species dis- 
tinctly belongs to the genus Pseudoxomysis 
in the characters of the maxilla, labrum and 
third to eighth thoracic endopods. From the 
other species of the genus, P. caudaensis, 
the new species is distinguished as shown 
in Table 1. 


Acknowledgments 


I would like to thank Dr. Mulyadi, Di- 
vision of Zoology, Research and Develop- 
ment Center for Biology, Indonesian Insti- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


tute of Science, for allowing me to examine 
the Pseudoxomysis caudaensis specimens. 


Literature Cited 


Liu R., & S. Wang. 1986. Studies on Mysinae (Crus- 
tacea Mysidacea) of the northern South China 
Sea.—Studia Marina Sinica 26:159—202. (in 
Chinese with English abstract.) 

Nouvel, H. 1973. Pseudoxomysis caudaensis n. gen., 
Nn. sp., crustacé mysidacé (Leptomysini) de la 
Mer de Chine méridionale.—Bulletin de la So- 
ciété d’ Histoire naturelle de Toulouse 109:131— 
141. 

Pillai, N. K. 1963. On a new mysid from the inshore 
waters of the Kerala coast.—Journal of the Ma- 
rine Biological Association of India 5:248—262. 

. 1964. Report on the Mysidacea in the collec- 

tions of the Central Marine Fisheries Research 

Institute, Mamdapam Camp, South India—Part 

I.—Journal of the Marine Biological Associa- 

tion of India 6:1—41. 

. 1973. Mysidacea of the Indian Ocean. Hand- 

book to the International Zooplankton Collec- 

tions, vol. 4, Papers on the Zooplankton Col- 
lections of the TOE, 125 pp. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(4):897—906. 2001. 


Redescription of Lepeophtheirus marginatus Bere, 1936 (Copepoda: 
Caligidae) and relegation of L. christianensis Wilson, 1944 and 
L. orbicularis Shino, 1960 as its synonyms 


Ju-shey Ho and Samuel G6mez 


(J-SH) Department of Biological Sciences, California State University, 
Long Beach, California, 90840-3702, U.S.A; 
(SG) Instituto de Ciencias del Mar y Limnologia, Unidad Académia Mazatlan, 
Joel Montes Camarena s/n, Ap. Postal 811, C.P. 82040, Mazatlan, Sinaloa, México 


Abstract.—Lepeophtheirus marginatus Bere, 1936 is redescribed based on 
examination of type specimens. Studying the type materials of L. christianensis 
Wilson, 1944 and L. orbicularis Shiino, 1965 from sea catfish (Galeichthyes 
sp.) collected in the Gulf of Mexico and Peru, respectively, led us to recom- 
mend relegating both of them as synonyms of L. marginatus Bere. 


In their report on a new species of Le- 
peophtheirus parasitic on the bullseye puff- 
er (Sphoeroides annulatus Jenyns) taken off 
Sinaloa, Mexico, Ho et al. (in press) sus- 
pected that L. christianensis reported by 
Wilson (1944) may be synonymous with L. 
marginatus Bere, 1936, because these nom- 
inal species resemble one another and prob- 
ably were reported from the same species 
of sea catfish (Galeichthys sp.) caught at the 
same locality (off Pass Christian, Mississip- 
pi). Therefore, type specimens of both spe- 
cies deposited at the National Museum of 
Natural History, Smithsonian Institution in 
Washington, D.C. were examined. Unex- 
pectedly, in the process of reviewing the 
species of Lepeophtheirus reported from the 
sea catfish, it was discovered that not only 
is L. christianensis synonymous with L. 
marginatus, but also is L. orbicularis Shi- 
ino, 1965. 

Based on these studies we herein rede- 
scribe L. marginatus and then discuss the 
establishment of L. christianensis and L. or- 
bicularis as junior synonyms of L. margin- 
atus. 


Materials and Methods 


The specimens kept in 70% ethanol in 
the collections of National Museum of Nat- 


ural History, Smithsonian Institution in 
Washington, D.C. and Faculty of Biore- 
sources, Mie University in Tsu, Japan were 
cleared in lactic acid for 1 hr before ex- 
amination. Two specimens (one female and 
one male) of L. marginatus from USNM 
79163 were dissected in a drop of glycerin 
and the removed body parts and append- 
ages were mounted on slides using glycerin 
as mounting medium and examined using a 
compound microscope. All drawings were 
made with the aid of a camera lucida. In 
this report a full description is given of the 
female and only those parts and appendages 
showing sexual dimorphism are given of 
the male. 


Lepeophtheirus marginatus Bere, 1936 
Figs. 1-6 


Material examined.—Type-material 1 9 
and 1 6 (USNM 69860) from “outside 
skin” of Arius felis (Linnaeus) (= Galei- 
chthys felis) collected at Englewood, Flor- 
ida and another collection containing 3 & 
2,2 6 6, and 1 young 2 (USNM 79163) 
from Lemon Bay, Florida (no host desig- 
nated on label). 

Female.—Body (Fig. 1A) 3.12 (2.94— 
3.44) mm long, excluding setae on caudal 


898 


rami. Cephalothoracic shield nearly as long 
(1.59 mm) as wide (1.43 mm), excluding 
marginal hyaline membranes. Fourth pedi- 
ger distinctly wider (373 wm) than long 
(193 wm). Genital complex (Fig. 1B) slight- 
ly longer than wide (942 X 878 wm) and 
scattered with setules in central part of dor- 
sal surface and posteroventral surface near 
leg 5; median portion of anterior margin 
protruded to form a short stem that connects 
to 4th pediger, posteroventral portion be- 
tween 5th legs with or without shield-like 
cement plate (originating from a male with 
whom female mated) holding spermato- 
phores. Abdomen (Fig. 1B) not clearly sep- 
arated from genital complex, distinctly wid- 
er than long (210 X 388 jm) and with con- 
vex sides. Caudal ramus (Fig. 1A, B) small, 
wider than long (51 X 70 wm), carrying 3 
short (mediodorsal seta broken off on both 
rami in dissected specimen) and 3 long plu- 
mose setae (broken off on both rami in dis- 
sected specimen). Egg sacs (not shown in 
Fig. 1A) longer than half body length. 
Frontal plate (Fig. 1A) with 1 setule on 
anterior margin near midline. Antennule 
(Fig. 2A) 2-segmented; proximal segment 
with 27 plumose setae on anterodistal sur- 
face, distal segment 2.69 times longer than 
wide, with isolated seta about midway on 
posterior margin and 11 setae plus 2 aesth- 
etascs on distal margin. Antenna (Fig. 2B) 
3-segmented; proximal segment smallest, 
with bluntly pointed postero-medial pro- 
cess; 2nd segment robust, with ventral cor- 
rugated pad near tip; distal segment with 
sharply pointed, bent tip, with small seta in 
proximal region and slender seta in middle 
region. Postantennal process (Fig. 2B) com- 
prising massive, globular base with 2 setu- 
le-bearing papillae and broadly rounded 
shaft. A spherical outgrowth located be- 
tween bases of antenna and postantennal 
process. Mandible (Fig. 2C) with 4 sec- 
tions, bearing 12 teeth on medial margin of 
distal blade. Maxillule (Fig. 2B) consisting 
of long, slender, pointed process and papilla 
with 3 unequal setae. Maxilla (Fig. 2D) 2- 
segmented; proximal segment (lacertus) 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


large, with transverse ridge across middle 
region and cuneiform process on medial 
surface; slender, distal segment (brachium) 
carrying small subterminal process on outer 
edge and 2 unequal elements (calamus and 
canna) terminally and subterminally. Max- 
illiped (Fig. 2E) 2-segmented; proximal 
segment (corpus) largest, with basal flange 
on medial surface; distal segments sharply 
pointed claw carrying small knob in prox- 
imal region and simple, medial seta in mid- 
dle region. Sternal furca on dissected spec- 
imen (Fig. 2F) with bluntly tipped tines 
bearing marginal hyaline membranes, but 
tines of other specimens appears spatula- 
like as shown in Fig. 6C. 

Armature on rami of legs 1—4 as follows 
(Roman numerals indicating spines and Ar- 
abic numerals, setae): 


Exopod Endopod 
Leg 1 1-0; LI, I, 3 (vestigial) 
ese I-1; I-1; Il, 5 0-1; 0-2; 6 
Leg 3 I-1; I, Il, 5 0-1; 6 
Leg 4 1-0; I-O; II (absent) 


Leg 1 (Fig. 3A) protopod with plumose 
outer seta and another plumose inner seta; 
endopod short, bluntly tipped pinnate pro- 
cess; first segment of exopod with row of 
setules on posterior edge and short spini- 
form seta at anterior-distal corner; middle 2 
of 4 terminal elements on last segment of 
exopod with accessory process, 3 terminal 
elements bearing crescent membrane ter- 
minally on both sides assuming spoon-like 
structure, and 3 posterior plumose setae 
short. Leg 2 (Fig. 3B) coxa small, with 
large plumose seta on posterior edge; basis 
with small, naked outer seta; both lateral 
and medial edge of protopod with large 
marginal membranous fringe, proximal lat- 
eral spines on third segment of exopod 
semipinnate with naked anterior side, next 
spine pinnate and appearing setiform. Leg 
3 (Fig. 4A) protopod with large lateral and 
posterior marginal membranous fringe in 
addition to lateral and medial plumose se- 
tae. Both rami 2-segmented; proximal seg- 


VOLUME 114, NUMBER 4 


Fig. 1. 
ventral. 


ment of exopod round, with marginal mem- 
branous fringe membrane and slender seti- 
form process; second and terminal seg- 
ments incompletely fused, with 2 setules 
proximal to lateral spines on terminal seg- 
ment. Leg 4 (Fig. 4B) protopod with short, 
naked distal-lateral seta; pectens on exopod 
segments at insertion of each lateral spine 
(Fig. 4C). 

Leg 5 (Fig. 4D) represented by pinnate 


899 


\ ma ie Ne 


me SP NS 2) So et Nk 
we Gf (yy aX K S. 


ee lain 8 |500 pm 


Lepeophtheirus marginatus Bere, 1936. Female. A, habitus, dorsal. B, genital complex and abdomen, 


seta and small process tipped with 3 plu- 
mose setae. 

Male.—Body (Fig. 5A) 2.27 (1.90—2.48) 
mm long, excluding setae on caudal rami. 
Cephalothoracic shield about as long (1.19 
mm) as wide (1.13 mm), excluding margin- 
al hyaline membranes. Genital complex 
(Fig. 5C) longer than wide, 916 X 786 wm, 
sparsely covered with setules on dorsal sur- 
face and lateral margins; ventral surface 


900 


ABDE 
100m 


path LA a Lia 
100m 


Cc 
400um 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Lepeophtheirus marginatus Bere, 1936. Female. A, antennule (not showing all setae), dorsal. B, 
antenna, postantennal process, and maxillule; ventral. C, mandible. D, maxilla. E, maxilliped. EF sternal furca. 


showing two posterolateral lobes bearing 
leg 6 at their tip. Abdomen (Fig. 5C) short 
and slightly wider (488 pm) than long (461 
wm). Caudal ramus (Fig. 5B) wider (68 
wm) than long (50 wm). Antenna (Al in 
Fig. 6A) 3-segmented; proximal segment 
with large corrugated pad; middle segment 
largest, robust and armed with 2 corrugated 
pads (Fig. 6B); terminal segment prehen- 
sile, quadrifurcate distally and armed with 
2 spiniform setae. Corrugated pad on ster- 
num (A3 in Fig. 6A) posterior and medial 


to maxillule (A2 in Fig. 6A). Dentiform 
process of maxillule with small protuber- 
ance near base (A2 in Fig. 6A). Corpus of 
maxilliped (Fig. 6D) with 2 transverse ridg- 
es near base; shaft bearing subspherical 
protuberance basally and spiniform seta dis- 
tally; claw with denticulate medial protru- 
sion near base (see insert drawing in Fig. 
6D). Sternal furca (Fig. 6C) with broad, 
spatula-like tines. Leg 5 (Fig. 5C) consist- 
ing of small plumose seta and papilla tipped 
with 1 simple and 2 plumose setae located 


VOLUME 114, NUMBER 4 


901 


Fig. 3. 


on lateral side of genital complex. Leg 6 
(Fig. SC) represented by 3 plumose setae at 
tip of ventral ridge on genital complex. 


Discussion 


The most distinguishing characteristics 
of L. marginatus are the possession of: a 
maxillule with simple (instead of bifid) den- 
tiform process (see Figs. 2B and 6A); a 2- 
segmented (instead of 3-segmented) exopod 
of leg 3 with slender setiform element (in- 
stead of robust, claw-like spine) on proxi- 
mal segment (see Fig. 4A); and a 3-seg- 
mented exopod of leg 4 with long (instead 
of minute), proximal, outer spine (see Fig. 
4B). Due to an incomplete original descrip- 
tion of L. marginatus and the lack of sup- 
plemental information in Causey (1955), 
the first two of these three unusual features 
of L. marginatus were unknown until now. 
Interestingly, these three features are also 


Lepeophtheirus marginatus Bere, 1936. Female. A, leg 1, anterior. B, leg 2, anterior. 


found in L. orbicularis Shiino and L. sim- 
plex Ho, Gomez & Fajer-Avila (2001). 
However, L. simplex is distinguishable from 
L. marginatus in the shape of the genital 
complex (oval in L. simplex) in both sexes, 
the structure of the sternal furca (with 
pointed and curved tines in L. simplex), the 
morphology of the terminal spines on the 
exopod of leg 1 (lacking crescent mem- 
brane in L. simplex), and the structure of 
the terminal claw of male antenna (bipartite 
and equipped with a tridentate medial pro- 
tuberance in L. simplex). 

One hundred and eight species of caligid 
copepods are currently classified in the ge- 
nus Lepeophtheirus. Among them, six spe- 
cies were reported from the sea catfishes 
(Ariidae). They are L. dissimulatus Wilson, 
1905; L. monacanthus Heller, 1865; L. un- 
ispinosus Pearse, 1952, L. christianensis, L. 
marginatus and L. orbicularis. Lepeo- 


902 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


we 
{ 7 
~ f 
<— emcee COO tower opener 


Meee 


heat 


Sx 


4 
So it bar 


D 


Fig. 4. Lepeophtheirus marginatus Bere, 1936. Female. A, leg 3, ventral. B, leg 4, anterior. C, terminal part 
of leg 4 exopod. D, leg 5. 


VOLUME 114, NUMBER 4 


903 


Fig. 5. 
complex and abdomen, ventral. 


phtheirus dissimulatus differs from L. mar- 
ginatus in the structure of leg 3, and L. 
monacanthus and L. unispinosus are distin- 
guishable easily from L. marginatus by the 
terminal armature on the exopod of leg 1. 
Thus, only L. christianensis and L. orbicu- 
laris remain to be scrutinized further. 

In her original report of L. marginatus, 
Bere (1936: 587) stated: ““A single male 
and female have been selected for the types 
of the new species with U.S.N.M. No. 
60548.” Curiously, Wilson (1944) listed the 
same catalogue number for his type mate- 
rial of L. christianensis. He (Wilson 1944: 


Lepeophtheirus marginatus Bere, 1936. Male. A, habitus, dorsal. B, caudal ramus, dorsal. C, genital 


533) stated: ““The female holotype and male 
allotype are U.S.N.M. No. 60548.” The 
type collections kept at Smithsonian Insti- 
tution show that the catalogue number of 
USNM 60548 is for the type lot of L. chris- 
tianensis and that of L. marginatus 1s 
USNM 69860. Strangely, although L. chris- 
tianensis was published eight years later 
than L. marginatus it is assigned a smaller 
(i.e., earlier) catalogue number. 

Another curiosity about Wilson’s report 
of L. christianensis concerns the number of 
specimens he collected, examined, and de- 
posited in the National Museum of Natural 


904 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Lepeophtheirus marginatus Bere, 1936. Male. A, antenna (A1), maxillule (A2), and corrugated pad 
(A3), ventral. B, middle and terminal segments of antenna, anterior. C, sternal furca. D, maxilliped, anterior. 


VOLUME 114, NUMBER 4 


History. Wilson (1944: 533) stated that, 
“Thirty specimens, including both sexes, 
were obtained from a sea catfish (Galei- 
chthys sp.) at Pass Christian, Miss.,”’ but 
museum records list a total of 32 specimens 
kept in two lots, two (one holotype female 
and one allotype male) in a lot carrying the 
catalogue number of USNM 60548 and 30 
(18 adult females, nine adult males, one 
pair in amplexus, and one juvenile female) 
in another lot identified as USNM 60549. 

Of the three distinguishing characteristics 
enumerated above for L. marginatus, the 
segmentation of the exopod of leg 3 and the 
armature on its proximal segment are the 
most remarkable feature of this species. In 
the original description of L. marginatus, 
Bere (1936: 588) stated: ““The second and 
third legs are of the usual type”’ and thus 
omitted the illustrations of these two ap- 
pendages. On the other hand, in Wilson’s 
(1944: 534) report of L. christianensis, 
these two appendages were neither men- 
tioned nor illustrated. As shown in Fig. 4A, 
leg 3 in L. marginatus is not “of the usual 
type.” It is rather unusual in bearing a 2- 
segmented exopod carrying a setiform ele- 
ment on the proximal segment. These un- 
usual features are found in all 32 specimens 
of L. christianensis deposited at the Nation- 
al Museum of Natural History. A check of 
their other features revealed that, indeed, all 
32 specimens are compatible with our re- 
description of L. marginatus. Accordingly, 
we recommend relegating L. christianensis 
Wilson, 1944 as a junior synonym of L. 
marginatus Bere, 1936. 

When Shiino (1965) reported L. orbicu- 
laris, nO comments on or comparison with 
L. marginatus were made. However, after 
observing the morphology of L. marginatus 
and noting its close resemblance to L. or- 
bicularis, we felt it was necessary to ex- 
amine the type material of the latter, which 
was kept in the Faculty of Bioresources at 
Mie University in Tsu, Japan. 

The type material of L. orbicularis (S- 
577) is kept in three vials marked 1, 2, and 
3, with each of them containing one, nine, 


905 


and five specimens, respectively. In his 
original description of L. orbicularis, Shi- 
ino (1965: 447) stated ““No. 577. Ten fe- 
males and five males. The largest female is 
selected as holotype.” But, close examina- 
tion of each specimen in a drop of lactic 
acid disclosed that while specimens in vial 
1 (containing holotype @) and vial 2 (con- 
taining 5 2 2 and 4 ¢ 3G) agree with Shi- 
ino’s (1965) description of L. orbicularis, 
the five specimens (4 2 @ and 1 d) kept 
in vial 3 are not. They differ from Shiino’s 
original description in the shape of the ab- 
domen (with straight sides) and structures 
on leg 3 (with longer velum) and leg 4 
(with slender exopod). At this time, those 
five specimens kept in vial 3 should be ex- 
cluded from L. orbicularis. 

Shiino’s (1965) description of L. orbi- 
cularis 1s clear and no supplemental infor- 
mation from our reexamination of the type 
material is significant to this study, except 
for the segmentation of the exopod of leg 
3, which is clearly 2-segmented as in L. 
marginatus. Although some minor differ- 
ences were detected in the fine structures of 
the maxillule (length of the dentiform pro- 
cess), maxilla (length of the subterminal 
process and ornamentation on calamus and 
canna), leg 2 (ornamentation on the two 
outer spines on the terminal segment of the 
exopod), and leg 4 (relative length of the 
lateral spine on the proximal segment of the 
exopod), these differences are considered 
geographical variation, because Shiino’s 
(1965) material came from Peru, a different 
locality from Bere’s (1936). Hence, we rec- 
ommend that the name L. orbicularis Shi- 
ino, 1960 be relegated as junior synonym 
of L. marginatus Bere, 1936. 


Acknowledgments 


We thank Chad Walter for arranging the 
loan of type-materials of L. marginatus and 
L. christianensis from the collection of the 
National Museum of Natural History, Smith- 
sonian Institution, and Izawa Kunihiko for 
the visit by one of us (JSH) to Mie Univer- 


906 


sity in Tsu, Japan to examine the type-ma- 
terial of L. orbicularis at that institution. Z. 
Kabata and two other reviewers are ac- 
knowledged for their comments and sugges- 
tions for the improvement of this paper. The 
completion of this manuscript was aided by 
a fellowship from the JISTEC (Japan Inter- 
national Science and Technology Exchange 
Center) and a grant from the Paramitas 
Foundation to the senior author (JSH). 


Literature Cited 


Bere, R. 1936. Parasitic copepods from Gulf of Mex- 
ico fish—The American Midland Naturalist 
17(3):577-625. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Causey, D. 1955. Parasitic Copepoda from Gulf of 
Mexico fish.—Occasional Papers of the Marine 
Laboratory 9:1—19. 

Ho, J. S., G. S. Noguera, & E. Fajer-Avila. 2001. Le- 
peophtheirus simplex, a new species of copepod 
(Siphonostomatoida: Caligidae) parasitic on 
“botete”’ (bullseye puffer, Sphoeroides annula- 
tus) in Sinaloa, Mexico.—Folia Parasitologica 
(in press). 

Shiino, S. M. 1965. Parasitic copepods of the eastern 
Pacific fishes. 8. Lepeophtheirus.—Report of 
Faculty of Fisheries, Prefectural University of 
Mie 5(2):441—454. 

Wilson, C. B. 1944. Parasitic copepods in the United 
States National Museum.—Proceedings of the 
United States National Museum 94(3177):529— 
582. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(4):907-917. 2001. 


Observations on Cumacea (Malacostraca: Peracarida) from Antarctic 
and subantarctic waters. I. Ekleptostylis debroyeri (Diastylidae), a new 


species from waters off the Antarctic Peninsula 


Magdalena Blazewicz-Paszkowycz and Richard W. Heard 


(MB-P) University of L6dz, Laboratory of Polar Biology and Oceanobiology, Pomorska 141, 


Lodz 90-237, Poland; 


(RWH) Department of Coastal Sciences, University of Southern Mississippi, Gulf Coast Research 


Laboratory Campus, Ocean Springs, Mississippi 39566-7000, U.S.A. 


Abstract.—Eighteen species of diastylid cumaceans have been described 
from Antarctic and subantarctic waters. A nineteenth species, Ekleptostylis 
debroyeri, n. sp., was collected in depths of 83 to 530 m off the Antarctic 
Peninsula. The new species appears to be closely allied to Ekleptostylis heardi 
McLelland & Meyer, 1998; E. inornata (Hale, 1937), n. comb.; E. pseudoi- 
nornata (Ledoyer, 1977); and E. vemae (Bacescu-Mester, 1967), all of which 
are known from subantarctic and Antarctic waters. Ekleptostylis debroyeri is 
distinguished from these species by a combination of characters including the 
size of the anterior-most pair of lateral spiniform setae of the telson and the 
proximal article of the uropodal endopod subequal to the combined length of 
the two distal most articles. The genera Diastylis Say, 1818, Ekleptostylis 
Stebbing, 1912, and Leptostylis G. O. Sars, 1869 have morphological features 
that appear to be transitional, especially among the subadults and females. 
The generic status of many species within these genera can only be deter- 
mined with certainty by the morphology (length of antenna, structure of the 
second pereopod, ornamentation of telson) of the terminal male stage. Al- 
though the mature male of EF. debroyeri is unknown, it is tentatively placed 
in Ekleptostylis based on the similarity of the female to other known species 


of the genus. 


This report represents the first in a series 
on the distribution, taxonomy, and system- 
atics of Cumacea from the Antarctic and 
subantarctic waters in depths ranging from 
3 to over 5000 m. Members of the cosmo- 
politan family Diastylidae are found 
throughout the world’s oceans and seas; 
however, most of the described species 
come from depths of greater than 1000 m 
(Day 1980, Jones 1969). In Antarctic and 
subantarctic waters (south of 50°S) diastyl- 
ids are presently represented by 18 nominal 
species and subspecies belonging to four 
genera. Table | lists these currently known 
diastylids. 

An examination of Antarctic and subant- 


arctic Cumacea from the collections in the 
Smithsonian Institution and Antarctic Bi- 
ology Program of the University of Lodz 
revealed an undescribed species tentatively 
referable to the genus Ekleptostylis Steb- 
bing, 1912. The description of the new spe- 
cies, which comes from waters in the vicin- 
ity of the Antarctic Peninsula, is the subject 
of this report. 

Type material has been deposited in the 
collections of the National Museum of Nat- 
ural History (USNM) and University of 
L6dz. Measurements are in mm unless oth- 
erwise stated and total body length (TL) is 
measured from the tip of pseudorostrum to 
tip of telson. 


908 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Type locality and depth of species and subspecies of the family Diastylidae presently known from 


Antarctic and subantarctic waters south of 50°E. 


Species Type Locality Depth (m) 
Diastylis anderssoni anderssoni Zimmer, 1907 South Georgia 64-385 
Diastylis anderssoni aramata Ledoyer, 1993 Weddell Sea (Halley Bay?)* 270-810 
Diastylis corniculata Hale, 1937 66°45’S, 62°03’E 218 
Diastylis enigmatica Ledoyer, 1993 75°14.2'S, 26°59.4’E 270-280 
Diastylis galeronae Ledoyer, 1993 74°32.2'S, 29°18.7'E 1165-1223 
Diastylis horrida Sars, 1887 Kerguelen Islands 20-540 
Ekleptosylis inornata (Hale, 1937), n. comb. 66°45'S, 62°03'E 150 
Diastylis mawsoni Calman, 1918 66°55'S, 145°06’E 200-582 
Diastylis zimmeri Ledoyer, 1977 Kerguelen Islands 18—90 
Diastylis annulata (Zimmer, 1902) South Georgia 4-355 
Diastylis diaphanes Zimmer, 1907 66°02’S, 89°03'E 385 
Diastylis goekei Roccatagliata & Heard, 1992 King George Island 2-399 
Ekleptostylis debroyeri, n. sp. South Shetlands, King George Is. 90—400 
Ekleptostylis heardi McLelland & Meyer, 1998 Argentina (54°04’S, 58°46’ W)*** 119 


Ekleptostylis pseudinornata (Ledoyer, 1977) 
Ekleptostylis vemae (Bacescu-Mester, 1967)** 
Holostylis helleri (Zimmer, 1907) 

Leptostylis antipa Zimmer, 1907 

Leptostylis crassidauda Zimmer, 1907 
Makrokylindrus baceskei Lomakina, 1968 
Makrokylindrus inscriptus Jones, 1971 


* No specific type locality designated. 


Kerguelen Islands 90 


Argentina (41°16’S, 60°03’W) 70-665 
South Georgia 12-640 
South Georgia 12-310 
66°02’S, 89°03’E 123-385 
64°03'W, 161°59.2'E 2938 
75°15'S, 166°15'E 808 


** See Roccatagliata & Miihlenhardt-Siegel (2000) for records from <50°S. 
*** See remarks in this report concerning a probable error for the original type-locality designation. 


Ekleptostylis debroyeri, new species 
Figs. 1-3 


Material examined.—Holotype, adult @ 
with developed oostegites, body length 10 
mm; USNM 1000896, Admiralty Bay 
(King George Island, South Shetland Is- 
lands, Antarctic), depth 290 m, transect 
Thomas Point—Urbanek Crag (sample no. 
OC-321), 17 Mar 1981, dredge, R/V Pro- 
fesor Siedlecki, BIOMASS-FIBEX Expe- 
dition (leg. Jazdzewski and Kittel).—Para- 
types (1d, 2 2 2), same collection data as 
holotype, Laboratory of Polar Biology and 
Oceanobiology, University of L6dz, Po- 
land. 

Other material examined.—Admiralty 
Bay (King George Island, South Shetlands): 
1 2, OC-324, 19 Mar 1981, depth 430 m; 
1 manca, OC-477, 11 May 1985, depth 211 
m, 1 manca, OC-487, 4 Sep 1985, depth 
162 m, 3 2 2, OC-733, 7 Dec 1988, depth 
400-530 m, 2 mancas OC-354, 13 Apr 


1983, depth 70 m; 1 2, 5 mancas, OC-517, 
30 Oct 1985, depth 212 m; 1 manca, OC- 
353, 13 Apr 1983, depth 286 m; 2 6 5 OC- 
341, 23 Mar 1981, depth 400-530 m; 1 2, 
OC-726, 10 Oct 1988, depth 240—280 m; 2 
2 2, OC-341, 23 Mar 1981, depth 400— 
530 m; 1 °%, OC€-348) ole442s— 
58°16.7'W; 1 2, OC-347, 58°56.6'S— 
61°45.5'W, depth 260-285 m, 26 Mar 
1981.—R/V_ Polarstern: 50+ specimens, 
Cruise 42, ANT XIV/2/175, 63°18.90’S— 
58°41.70’W 23 Dec 1996, depth 491 m,.— 
R/V Hero: 1 2, Cr. 731/1847, 67°52.00’S— 
68°56.00'W, 26 Feb 1973, depth 185 m; 1 
subadult 6 Cr. 731/1823, 64°47.23'S— 
64°07.20'W; 20 Feb 1973, depth 90-110 
m.—R/V Eltanin: 5 2 2 Cruise 6, Sta. 439, 
63°50.10'S—62°35.90'W, 9 Jan 1963, depth 
128-165 m. 

Diagnosis.—Carapace approximately 
0.25 total body length, lacking spines, 
sparsely covered in fine hair-setae. Pseu- 


VOLUME 114, NUMBER 4 


Fig. 1. 
2, another aspect (setae on 2 proximal article not shown); E—I, pereopods 1—5 (exopod not illustrated for pereopod 
1); J, left uropod, telson last abdominal somite. Scales: 1 = 1.0 mm for A; 2 = 0.2 mm for B—D; 0.3 mm for 
E-J. 


dorostral lobes slightly downturned with 
dorsal margins straight. Antennal notch ab- 
sent in both sexes. Telson slightly longer 
than last abdominal segment, constricted 
posteriorly, bearing 2 pairs of lateral, setu- 
late spiniform setae with pair of simple, 
fine, dorsal-lateral setae between them; pos- 
terior tip with terminal pair of spiniform se- 
tae strongly developed. Carpus and propo- 
dus of pereopod | and 2 with short stout 


909 


Diastylis debroyeri, n. sp. (female) A, lateral view of adult; B, antenna 1; C, antenna 2; D, antenna 


distal setae. Uropods slender, more than 
twice as long as telson. 
Description.—Female (based on adults 
unless stated otherwise). Body length 9-11 
mm, sparsely covered with fine simple setae 
(Fig. 1A). Carapace approximately 1.5 as 
long as deep, twice as long as combined 
length of pereonites and approximately 4 
total body length. Pseudorostral lobes de- 
flected slightly downward, dorsal margins 


910 


straight. Distinct antennal notch not evi- 
dent. 

Abdomen subequal in length to cepha- 
lothorax; pleomeres increase successively 
in length until 5th; 6th subequal to 4th. 

Telson (Fig. 1J). Constricted posteriorly, 
length (excluding terminal spiniform setae) 
subequal to last abdominal segment; post- 
anal region subequal in length to preanal, 
bearing 2 pairs of spiniform (setulate) setae 
laterally, 1 pair of subdistal, dorsal-lateral 
fine simple setae between lateral spiniform 
pairs; tip of telson armed with pair of well- 
developed terminal spiniform setae. 

First antenna (Fig. 1B). Peduncle and fla- 
gellum having 3 and 5 articles, respectively. 
Peduncle article 1 stout, bearing | subdistal 
simple seta, 1 distal plumose seta, and clus- 
ter of numerous hair setae distoventrally; 
peduncle articles 2 and 3 attenuated, much 
narrower than article 1, each with large dis- 
tal simple seta. Flagellum with 5 articles; 
accessory flagellum minute, with 3 articles. 

Second antenna (Fig. 1C, D). Small, 
poorly developed, having 3 short articles on 
buttressed carina with plumose seta near at- 
tachment with article 1; article 1 wedge 
shaped with 1 simple spiniform seta; article 
2 asymmetrical, broadest distally, with long 
well-developed plumose setae on distal 
lobe; terminal article small, rounded distal- 
ly with single terminal plumose seta. 

Mandible (Fig. 2A, B). Slender (typical 
diastylid shape); left mandible bearing la- 
cinia mobilis and 9 spiniform setae, 4 den- 
tate and 5 finely serrate, between molar and 
incisor process; right mandible with | distal 
palmate comb seta and 9 finely serrate se- 
tae. 

Labium (Fig. 2C). As illustrated. 

Epignath (Fig. 2D). As illustrated. 

First maxilla (Fig. 2E). Outer endite with 
14 relatively slim setae of various shapes; 
inner endite with 5 setae (4 of different 
shapes and | accessory); palp with 2 distal 
setae of different lengths armed distally 
with microspinules. 

Second maxilla (Fig. 2F). Protopodite 
wide with numerous setae of different 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


shapes, outer endite with 6 spiniform setae; 
inner endite with 4 spiniform setae (3 blunt- 
ly serrate, 1 non-serrate with basal setule). 

First maxilliped (Fig. 2G, J). Siphon long 
with terminal lobes wide and relatively 
short, appearing fused (Fig. 2J). Endopodal 
articles stout and wide with numerous setae 
along inner margin, ischium reduced, car- 
pus with 2 long plumose setae distally (oth- 
er setation as illustrated), dactyl slender 
bearing 5 terminal or subterminal setae. 

Second maxilliped (Fig. 2H). Basis equal 
to combined length of merus and carpus, 
with 1 long sparsely plumose seta and 4 
small simple, needle-like setae distally; is- 
chium approximately 0.5 length of merus 
with 1 plumose seta on inner margin. Merus 
approximately length of carpus with plu- 
mose setae on middle of inner margin and 
subdistal on outer margin; carpus twice as 
long as propodus with numerous setae on 
inner margin and 2 subdistal plumose setae 
on outer margin; propodus with numerous 
setae on inner margin, | long, plumose seta 
medially and 1 subproximal long plumose 
seta; dactyl slender, tipped by 4 simple ter- 
minal and subterminal simple setae. 

Third maxilliped (Fig. 21). Endopod: Ba- 
sis approximately 1.5 longer than remaining 
articles combined; inner margin with 12 
plumose setae; distal process well-devel- 
oped with 3 stout, relatively short plumose 
setae (not reaching tip of propodus); ischi- 
um 0.5 length of merus with small plumose 
seta on inner margin; merus twice as long 
as ischium, inner margin with 3 small plu- 
mose setae, outer with | distal plumose 
seta; carpus, propodus and dactyl subequal; 
carpus with plumose setae along inner mar- 
gin, 1 distal seta on outer margin; propodus 
slender with simple setae distally on inner 
margin and | plumose on outer; dactyl with 
simple terminal and subterminal setae. Ex- 
opod: well-developed, excluding setae, 
nearly reaching to distal end of basis of en- 
dopod. 

First pereopod (Fig. 1E). [based on sub- 
adult 2] Attenuate, long with ischium, mer- 
us, carpus, propodus, and dactyl with few 


VOLUME 114, NUMBER 4 911 


| 


see 


Fig. 2. Diastylis debroyeri, n. sp. A-B, mandible; C, labium; D, epignath; E, maxilla 1; K maxilla 2, with 
detail (above) of outer and inner endite viewed from outer face; G, maxilliped 1, inner face (with detail of 
setation); H, maxilliped 2; I, maxilliped 3; J, terminal lobes of siphon (ex. maxilliped 1). Scales: 1 = 0.1 mm 
for A; 0.2 mm for B; 2 = 0.2 mm for C—E; 0.1 mm for F; 3 = 0.2 mm for G, J; 4 = 0.4 mm for H, I. 


912 


simple setae; basis less than 0.33 total 
length of appendage, ventral margin fringed 
with plumose setae; ischium and merus 
subequal; carpus elongate, subequal to pro- 
podus; dactyl about 0.33 length of propo- 
dus. Exopod well-developed excluding se- 
tae, reaching to carpus of endopod. 

Second pereopod (Fig. 1F). Basis sube- 
qual in- length to carpus; merus subequal to 
propodus; dactyl about 1.5 as long as pro- 
podus; each article with numerous simple 
setae with carpus and propodus also having 
stout spiniform setae distally. Exopod well- 
developed, as in first pereopod. 

Third pereopod (Fig. 1G). Basis slender, 
approximately 1.5 as long as remaining ar- 
ticles combined, with some simple setae; is- 
chium approximately 0.5 length of merus; 
carpus as long as ischium and merus com- 
bined, simple lateral setae present, 3 long 
simple setae on distal margin (extending 
past tip of dactyl); propodus as long as dac- 
tyl, with 1 distal seta; dactyl slightly longer 
than wide, tipped by 2 simple setae. Exopod 
greatly reduced, with 2 articles, distal arti- 
cle twice as long as proximal. 

Fourth pereopod (Fig. 1H). Similar to 
third, but stouter, basis as long as remaining 
articles combined; ischium and merus with 
simple seta. Exopod reduced, as in third pe- 
reopod. 

Fifth pereopod (Fig. 11). Similar to 
fourth, but basis shorter than remaining ar- 
ticles combined. Exopod lacking. 

Uropod (Fig. 1J). Slender, elongate. Pe- 
duncle over twice as long as telson with 
12-14 spiniform setae along inner margin; 
outer margin with some fine simple setae. 
Endopod with 3 articles; article 1 longer 
than 2 and 3 combined, bearing 3 spiniform 
setae along inner margin; articles 2 and 3 
subequal, article 2 with distal spiniform seta 
on inner margin, article 3 simple terminal 
and subterminal setae. Exopod with simple 
setae only, reaching just beyond article 2 of 
endopod; proximal article reduced, distal 
article with marginal setae along distal 0.75 
and 3 terminal setae. 

Subadult male, 


paratype (Fig. 3A). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Length 9.2 mm. Similar to female in exter- 
nal view, except for much greater devel- 
opment of exopods and broadening of basis 
on pereopods 3 and 4. 

Telson (Fig. 31). Having 3 pairs of lateral, 
setulate, spiniform setae (instead of 2 as in 
female). 

First antenna (Fig. 3B). Peduncle with 3 
stout articles decreasing in length distally, 
article 3 cubbed (stout, rounded); flagellum 
with 7 articles; accessory flagellum with 3 
articles, extending to mid-region of flagel- 
lum article 3. 

Second antenna (Fig. 3C). Flagellum 
about 0.5 body length. 

First pereopod (Fig. 3D). Basis longer 
than combined length of ischium, merus 
and carpus, with numerous simple setae; is- 
chium subequal to merus; carpus elongate; 
propodus and dactyl lost. 

Second pereopod (Fig. 3E). Basis broad 
subequal to combined length of ischium, 
merus, carpus and propodus; dactyl elon- 
gate, 3 times longer than propodus; each 
article with some simple setae; carpus and 
propodus with stout distal spiniform setae. 

Third pereopod (Fig. 3F). Basis broad, 
longer than remaining articles combined; is- 
chium approximately 0.5 of merus length 
with 2 distal seta; carpus as long as com- 
bined length of ischium and merus, with 
few lateral setae and 2 distal setae; propo- 
dus subequal to dactyl, with one terminal 
setae; dactyl with 2 terminal setae. Exopod 
well developed. 

Fourth pereopod (Fig. 3G) similar to 
third. 

Fifth pereopod (Fig. 3H) as in female. 

Uropod (Fig. 31). Similar to female, but 
with 16 lateral spiniform setae along inner 
margin of peduncle and endopod, and more 
simple setae on rami. 

Etymology.—This species is named in 
honor of Claude DeBroyer, Institut Royal 
Sciences Naturelles de Belgique, in recog- 
nition of his many significant contributions 
to carcinological research in the Antarctic. 

Remarks.—Ekleptostylis debroyeri is 
similar to Ekleptostylis heardi McLelland & 


VOLUME 114, NUMBER 4 913 


Fig. 3. Diastylis debroyeri, n. sp. (subadult male) A, lateral view of adult; B, antenna 1; C, antenna 2 of 
subadult; D—H, pereopods 1—5 (propodus and dactyl missing on pereopod 1, exopod illustrated only for pereopod 
2); I, uropods, telson, and last abdominal somite. Scales: 1 = 1.0 mm for A, C; 2 = 0.2 mm for B—H; 0.3 mm 
for I. 


914 


Meyer, 1998; E. pseudoinornata, (Ledoyer, 
1977); E. vemae Bacescu-Mester, 1967; and 
Diastylis inornata Hale, 1937, all of which 
are known from subantarctic waters. Based 
on the descriptions and illustrations of D. 
inornata given by Hale (1937), Ledoyer 
(1977) and examination of the type material 
from the Kerguelen Islands, E. debroyeri 
can be distinguished from it by: the proxi- 
mal article of the uropodal endopod sube- 
qual to the combined length of the two dis- 
tal articles; differences in the setation on the 
third maxilliped; carpus of pereopod 2 
armed with short, stout, distally located spi- 
niform setae; the carpal setae of the third, 
fourth, and fifth pereopods extending be- 
yond those of dactyl; more slender uropods; 
telson half or less as long as uropodal pe- 
duncle with an additional pair of delicate 
setae. Ekleptostylis debroyeri is further dis- 
tinguished from D. inornata by the rela- 
tively straight dorsal margin of its pseudo- 
rostrum. 

Ekleptostylis pseudinornata, recently 
transferred from the genus Diastylis Say, 
1818 by Roccatagliata & Miihlenhardt-Sie- 
gel (2000), is distinguished both from E. 
debroyeri and D. inornata by possessing an 
antennal notch. Ledoyer (1977) described 
D. pseudoinornata based on two speci- 
mens, an adult (marsupial) female holotype 
bearing oostegites and a male paratype, col- 
lected in Kerguelen Islands. He distin- 
guished his female specimen of E. pseu- 
doinornata from D. inornata primarily on 
the presence of a pair of anterior-dorsal pro- 
jections on the first two pereonites, the 
presence of an antennal notch, and the den- 
tition of the anteroventral margin of the car- 
apace. Study of material attributable to D. 
inornata in the Institut Royal Sciences Na- 
turelles de Belgique, Brussels, revealed that 
adult females with developed oostegites 
also have these same dorsal projections on 
the first two pereonites, but those of smaller 
subadult, preincubatory females lack these 
tubercles. We reexamined the holotype and 
paratype of D. inornata, which are both 
subadult females, and found no indication 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


of anterodorsal projections observed in the 
adult females of E. pseudoinornata, E. 
heardi, and those in the Brussels collection, 
which were referred to D. inornata. 

Although no adult males or incubatory 
females are known for D. inornata, based 
on the features of the carapace, uropods, 
and telson, we believe that it represents an- 
other member of the genus Ekleptostylis 
and transfer D. inornata to the genus Ek- 
leptostylis here. In addition to the descrip- 
tions of the antenna, second pereopod, and 
telson for the terminal male of Ekleptestylis 
inornata (Hale, 1937), n. comb, detailed de- 
scriptions and illustrations of the mouth 
parts of adult males and females with fully 
developed oostegites collected from the 
Kerguelen Islands are needed. 

The adult female of E. debroyeri can be 
distinguished from those of E. walkeri (Cal- 
man, 1907) (type species), E. heardi, and 
E. vemae, by a having a telson bearing only 
two pairs of lateral spiniform setae plus a 
pair of characteristic simple setae (Figs. 1J, 
31). The telson of adult female E. walkeri, 
E. vemae, and E. heardi are armed with 
10+, 3—4 and 4—5 pairs of lateral spiniform 
setae, respectively. 

The adult female of E. debroyeri is fur- 
ther distinguished from that of E. vemae by 
having: a maxilliped 3 without an inner dis- 
tal spine (or spines) on basis; different pro- 
portions for the articles in pereopods 1 and 
2; a distinctly larger and longer (10 mm) 
body; a distinctly longer pair of terminal 
setae on the telson; the peduncle of the uro- 
pod two or more times longer than the tel- 
son; first article of the uropodal endopod 
distinctly longer than both two distal arti- 
cles combined (not subequal as in E. ve- 
mae). 

Besides the larger number of lateral spi- 
niform setae, Ekleptostylis debroyeri can be 
separated from E. heardi by its relatively 
shorter telson. In E. heardi the telson is 
over 2/3 the length of the uropodal pedun- 
cle. 

There is confusion concerning the type 


VOLUME 114, NUMBER 4 


locality of E. heardi and the depth and lo- 
cality data for the type material presented 
by McLelland & Meyer (1998) appears to 
be in error. Based on the written label, some 
specimens of FE. heardi, including the type 
material, came from the abyssal depth of 
3590 m. There is strong circumstantial ev- 
idence that Eltanin Station ‘°363”’ (Cruise 
6) was an apparent mislabeling of one of 
the fractions of the large “rock dredge” 
sample taken at Eltanin Station 343. Station 
363 was a Phleger corer sample taken at a 
depth of 3477-3590 m. In addition to Ek- 
leptostylis heardi, three other diastylid spe- 
cies, Diastylis hammoniae Zimmer, 1902, 
Diastylis planifrons Calman, 1912, and 
Diastyloides goekei Roccatagliata & Heard, 
1992 were also present in the vial labeled 
“Station 363.” All three species were pre- 
viously known from Antarctic and subant- 
arctic waters in depths of less than 400 m 
(Roccatagliata & Heard 1992, Calman 
1912). When received from the University 
of Southern California where the Eltanin 
samples were sorted, the vials labeled Sta- 
tions 343 & 363 were placed together in a 
large vial (Heard, per. obs.), further sup- 
porting our supposition of a mislabeling er- 
ror. 

Due to this apparent mislabeling during 
the sorting process, McLelland & Meyer 
(1998) listed (p. 279) their type material of 
Ekleptostylis heardi as coming from “Sta. 
363” at a depth of 3590 m.”’ We recom- 
mend that all their material should now be 
referred to a single locality, Eltanin Station 
343 (Cruise 6), and that the type locality of 
E. heardi be changed to accommodate this 
apparent error. 

Discussion.—Subadult specimens that 
we attribute to E. debroyeri, appear to be 
nearly indistinguishable from the genus 
Leptostylis G. O. Sars, 1869, sensu lato. 
Day (1980) noted that the distinction be- 
tween Diastylis and Leptostylis cannot be 
determined without the adult males, since 
other generic characters appear to be too 
plastic. This situation also appears to apply 
to the females and subadults of Ekleptos- 


915 


tylis. As defined by Day (1980), the telson 
of Diastylis is usually longer than the last 
pleomere and has at least three pairs of lat- 
eral spiniform setae. In contrast, the telson 
of Leptostylis is short and supposedly never 
longer than the last pleomere (Day 1980). 
Another important diagnostic character at- 
tributed to Leptostylis is based on the length 
of second antenna of the adult male, so 
complete determination has not been pos- 
sible because the males for some species 
remain unknown. The descriptions of four 
species attributed to Leptostylis are based 
solely on subadult males. These species, 
Leptostylis faurei Day, 1980, L. gilli Day, 
1980, and L. menziesi Bacescu-Mester, 
1967, have a “‘cub-shaped”’ article 3 on sec- 
ond antenna and the flagellum of the second 
antenna barely reaches half the body length. 
According to the definition of Leptostylis 
sensu Stebbing (1912), this character would 
distinguish them from Diastylis. However, 
since their descriptions, as well as ours for 
E. debroyeri, are based on subadult males, 
there is a possibility that the terminal male 
forms might have considerably longer sec- 
ond antennae and thus affinities with the ge- 
nus Diastylis sensu Day 1980. Also, the tel- 
sons of these species are subequal, or even 
longer than sixth pleomere, and with the ex- 
ception of L. menziesi, they have 3 or more 
pairs of lateral spiniform setae on the post- 
anal region of the telson. These characters 
further suggest affinities to Diastylis sensu 
lato or possibly Ekleptostylis. 

The generic status of these three species 
will not be settled with certainty until adult 
males and females are available for study 
or until a more reliable set of characters can 
be established to separate Diastylis, Eklep- 
tostylis, and Leptostylis. The taxonomy of 
these three genera also requires designation 
of a type species for both Diastylis and Lep- 
tostylis. 

The taxonomic status of D. arenaria Say, 
1818, the type species and genotype for the 
family Diastylidae may be nearly resolved 
(see Gerken 1999). The type material for D. 
arenaria is poorly defined and no longer 


916 


extant (Holthuis 1969). The brief and vague 
species description was based on specimens 
collected from the east coast of the United 
States [““Coast of Georgia and Florida” 
(Say 1818:315)]. Diastylis arenaria may be 
conspecific with Oxyurostylis smithi Cal- 
man, 1912, the well-described type species 
of the genus Oxyurostylis Calman, 1912, 
which also was described from the shallow 
waters of the U. S. East Coast (Holthuis 
1969, Roccatagliata & Heard 1995). As 
suggested by Day (1980), Gerken (1999) 
petitioned the International Commission on 
Zoological Nomenclature to suppress D. ar- 
enaria as the type species in favor of the 
well established Diastylis rathkei (Kroyer 
1841). At this time her case (3078) is still 
pending. With regard to genus Lepfostylis, 
apparently Sars (1869) did not designate a 
type species for one of the four initial spe- 
cies assigned to the genus (see Day 1980: 
Di1@): 


Acknowledgments 


We are grateful to Krzysztof Jazdzewski 
and W. Kittel (University of Lodz) as well 
as Oliver Coleman (Museum fiir Naturkun- 
de, Berlin) for making the specimens avail- 
able for the study. We thank Krzysztof Jaz- 
dzewski and Sara LeCroy, John Foster, and 
Jerry McLelland, (Department of Coastal 
Sciences, University of Southern Mississip- 
pi [COA]) for their constructive comments 
on the earlier drafts of the manuscript. 
Dawne Hard (COA) kindly provided edi- 
torial assistance with the text and figures. 
We also wish to thank Cheryl Bright and 
William Moser (Department of Invertebrate 
Zoology, National Museum of Natural His- 
tory, Smithsonian Institution) for their help 
and support during the senior author’s stay 
in Washington, D.C. Wolfgang Zeidler 
(South Australian Museum) graciously 
loaned us the type material of D. inornata 
and Claude DeBroyer (Institut Royal Sci- 
ences Naturelles de Belgique) kindly made 
specimens attributable to D. inornata avail- 
able for study. We appreciate the helpful 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


suggestions and constructive criticisms 
made by three anonymous reviewers on 
earlier draft of the manuscript. This re- 
search was supported by a grant from the 
National Museum of Natural History, De- 
partment of Invertebrate Zoology, U.S. Ant- 
arctic Research Program funded by the Na- 
tional Science Foundation (Contract OPP- 
9509761). 


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Say, T. 1918. An account of the Crustacea of the Unit- 
ed States. Reprinting by J. Cramer, S-H. Service 
Agency, Inc., New York, 138 pp. 

Stebbing, T. R. R. 1912. South African Crustacea. Part 
6. The Sympoda.—Annals of the South African 
Museum 10:129—-176. 

Zimmer, C. 1902. Cumaceen.—Ergebnisse der Ham- 
burger Magalhaensische Sammelreise 2:1—18. 

. 1907. Neue Cumaceen aus der familien Dias- 

tylidae und Leuconidae von der Deutschen und 

der Schwedisch Sudpolar-Expedtion.—Zoolo- 

gischer Anzeiger 31:367—374. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(4):918—928. 2001. 


Probopyrus insularis, a new species (Isopoda: Bopyridae), a parasite 
of Macrobrachium faustinum (Saussure de, 1857) (Decapoda: 
Palaemonidae), with criteria to differentiate species of Probopyrus 


Ramiro Roman-Contreras and Roland Bourdon 


(RRC) Laboratorio de Parasitologia Marina, Instituto de Ciencias del Mar y Limnologia, 
Universidad Nacional Autonoma de México (UNAM). P.O. Box 70-305, Mexico 04510. E-mail: 
ramiror @ mar.icmyl.unam.mx 
(RB) 1 Impasse Corbiére, Roscoff 29680, France 


Abstract.—Probopyrus insularis n. sp. parasiting Macrobrachium faustinum 
(Saussure de, 1857) from the Caribbean Islands is mainly distinguished by the 
shape of the: pleotelson in both sexes; carina of the seventh legs in females; 
palp of the maxillipeds in females; and pleon in males. Characters used to 
distinguish species of Probopyrus Giard & Bonnier, 1888 are discussed. 


Two new American species of Probopy- 
rus recently have been described as part of 
an ongoing taxonomic study of these para- 
sitic isopods. Probopyrus pacificensis Ro- 
man-Contreras, 1993, and P. markhami Ro- 
man-Contreras, 1996, both were described 
from the west coast of Mexico. Here a new 
Probopyrus parasitizing Macrobrachium 
faustinum (Saussure de, 1857) from Jamai- 
ca, Puerto Rico, Guadeloupe and Cuba is 
described. Other American species new to 
science are expected as taxonomic research 
continues. 

Holthuis (1977) recorded M. faustinum 
parasitized by “bopyrids”’ from Cuba; and 
Kensley & Schotte (1989), and Bunkley- 
Williams & Williams (1998) have assigned 
M. faustinum as host of Probopyrus pan- 
dalicola (Packard, 1879). These records, 
however, might be a misidentification of the 
new species described here. Macrobrach- 
ium faustinum, the only host known for this 
new species, has been reported from the 
United States, Venezuela and the Antilles 
by Chace & Hobbs Jr. (1969), Holthuis & 
Provenzano (1970), Chace (1972), Dugger 
& Dobkin (1975), Pereira (1991), Fiévet 
(1998, 1999a, 1999b, 2000), Fiévet et al. 
(1996, 1999) and Bowles et al. (2000). The 
new isopod may be present in these areas. 


In a previous paper Romdan-Contreras 
(1993) summarized and proposed a series 
of diagnostic characters for the genus Pro- 
bopyrus. These characters are updated and 
applied to this description (Tables 1 and 2). 


Genus Probopyrus Giard & Bonnier, 1888 
Probopyrus insularis, new species 
Figs. 1-22 


Holotype female.-—USNM 120073. Al- 
lotype male USNM 306876, prepared for 
SEM; C. W. Hart Jr. coll., 7 Jan 1960, Trib- 
utary of Trunnels River, St. Mary Parish, 
Jamaica. 

Other material examined.—USNM 
235988: one young female, Rio Cruces, 
1500 m W of Sabana Grande, Puerto Rico. 
USNM 235990: one female, one male, 8 
May 1953, Rio Lajas, 3000 m S of Vega 
Alta, Puerto Rico. USNM 63309: one fe- 
male, one male, J. C. Welsh Jr. coll., Cuba. 
One female, one male, Serre coll., 1910, 
Cuba (in the Muséum National d’ Histoire 
Naturelle, Paris). Twenty one females and 
males, Y. Thérézien coll. and det. hosts, 2 
Dec 1977, Belle Eau, village de Il’ Habituée, 
Guadeloupe. Ten females, eight males, Y. 
Theérézien coll. and det. hosts, 30 Oct 1978, 
Lézarde estuary, Guadeloupe. Twenty nine 


VOLUME 114, NUMBER 4 


females, 26 males, E. Fiévet coll. and det. 
hosts, Dec 1996, Bananie River, Guade- 
loupe. Material from Guadeloupe deposited 
at collections of the authors. 

Type locality.—Tributary of Trunnels 
River, St. Mary Parish, Jamaica. 

Distribution.—Jamaica, Puerto Rico, 
Cuba, and Guadeloupe. 

Host for all specimens.—Macrobrach- 
ium faustinum. 

Habitat.—Freshwater. 

Description of holotype female.—Length 
10.3 mm, from border of first pereonite to 
tip of sixth pleonite; maximal width at third 
pereonite, 7 mm. Distortion angle 26°, out- 
line pyriform (Figs. 1, 2). Head almost as 
wide as long, front rounded, posterior bor- 
der deeply inserted into first pereonite, sur- 
face smooth. No well defined frontal lami- 
na, indicated by only a narrow line; antero- 
lateral corners of head produced into slight 
rounded tips. Antennula and antenna bis- 
egmented; antennula ovoid, 4—5 spines on 
distal tip; antennae smaller than antennulae, 
basal portion wider than antennulae, distal 
one subspheric-shape (Fig. 3); both anten- 
nae with small dispersed pectinate scales on 
surface. Maxilliped bisegmented, subovoid, 
anterolaterally convex, nonsetose palp 
formed by two digitiform processes (Fig. 
4). Three pairs of smooth, lamellar projec- 
tions on barbula; inner lateral projections 
lanceolate, outer ones slightly larger and 
wider, third one smaller than former and be- 
neath them; middle region rounded (Fig. 5); 
eyes absent. Pereonites dorsally distinct; 
border of pereonites 1—4 produced into dis- 
tinct dorsolateral bosses, coxal plates nar- 
row (Fig. 2). Oostegites completely sur- 
rounding but not enclosing marsupium; 
oostegite 1 relatively large, partially cov- 
ering head and anterior part of brood pouch; 
anterior segment transversely ovoid and 
concave, frontal border rounded and slight- 
ly undulate; distal segment digitiform, larg- 
er axis perpendicular to the former; digiti- 
form process on inner ridge, distal portion 
smooth, small spines on inner border; retic- 
ulated black pigmentation on surface, ex- 


919 


cept on distal portion (Fig. 6). Oostegites 
2—4 progressively longer; oostegites 3—4 on 
larger side unpigmented; fifth oostegites 
longer than others, slightly falcate, ending 
in blunt tip, densely setose along posterior 
margin, extending across posterior region 
of marsupium and overlapping each other. 
Pereopods with massive basipodite, dacty- 
lus hook shape (Fig. 7). Robust high carina 
on pereopods, either rectangular (P1—P2), 
convex (P3—P4) or subquadrangular (P5— 
P7) (Fig. 8). Pigmentated spots on basis of 
pereopods 1—5 on short side; basis of pe- 
reopods 6—7 on short side, and 1—7 on larg- 
er side, unpigmentated. Pleon 0.3 times 
body length, five dorsal and laterally sepa- 
rated pleonites; lateral plates lamellar, al- 
most rounded or slightly truncated. Pleotel- 
son bell-shaped, relatively small, bilobated, 
deep fissure in the middle, lobes unseparat- 
ed, tips shorter than the fifth lateral plates 
(Fig. 2). Five pairs of biramous foliate ple- 
opods completely covering ventral surface 
of pleon, each different in shape and size; 
exopods on large side enlarged, increasing 
in length from first to fifth, protruding no- 
tably from tips of pleon (Figs. 1, 2, 15); 
endopods decreasing in size from first to 
fifth; larger exopods on opposite side slight- 
ly rippled (Fig. 9); uropods absent. 
Description of male.—Length 1.84 mm, 
width 0.6 mm; body unpigmented. Head 
twice as wide as long, slightly depressed at 
middle (Figs. 10, 12, 13); posterior margin 
almost straight, anterolateral borders round- 
ed. Antennae different in size and shape; an- 
tennula three-segmented, bottle-shape, prox- 
imal portion globose, upper portion small, 
button shape, eight obscure small spines on 
tip. Antenna bisegmented, slightly smaller 
and thinner than antennula, cylindrical, with 
short proximal basis; stout short setae on tip 
of distal article. Both antennae with small 
pyramidal or rounded pectinate scales on 
surface (Fig. 11). Maxillipeds absent. No 
eyes. Pereon wider at second pereonite, 4—7 
progressively shorter; tips of pereonites 
subglobose, reflexed ventrally, small scales 
on ventral surface; midventral tubercles ab- 


920 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1—Morphological comparative characters for the American species of Probopyrus (Females). (Hosts: 
M. = Macrobrachium, P. = Palaemonetes, Pa. = Palaemon).* 


Characters 


Concavity on Cephalon: 
short side Body, Pigmentation on shape of the Cephalon: 
Species/host Body shape of body distal region pereomeres 2-4 posterior region frontal border 
P. insularis/ Pyriform Present Pointed On short Rounded, Convex 
M. faustinum side ovoid, or 
subtriangular 
P. bithynis/M. ohionis Pyriform Present Pointed On short Triangular Right 
(sic) side 
P. bithynis/M. olfersii 
P. markhami/ Ovoid or Absent Rounded On short Subpentagonal Right 
M. americanum pyriform side 
P. meeki/M. acanthurus Pyriform Slight Rounded Absent Ovoid Convex 
P. pacificensis/ Pyriform Present Pointed On short Ovoid Right 
M. tenellum side 
P. panamensis/ Pyriform Present Pointed On short Ovoid Convex 
M. acanthurus side 
P. floridensis/ Pyriform or Present or Pointed On short Ovoid or Right or 
P. paludosus ovoid absent side triangular acute 
P. pandalicolal Pyriform Slight Rounded On 2 sides Rounded Convex 
P. hiltonii 
P. pandalicola/P. pugio Ovoid Absent Rounded On 2 sides Rounded or Right or 
ovoid convex 
P. pandalicola/ Pyriform, Present or Pointed or On 2 sides Rounded or Right or 
P. vulgaris ovoid or absent rounded triangular convex 
rounded 
P. ringueleti/ Ovoid Absent Pointed Ovoid Acute 
P. argentinus 
P. palaemoni/ Ovoid Absent Pointed or On 2 sides Rounded or Right or 
Pa. pandaliformis rounded triangular convex 


4 Sources: see Table 2. 


sent. Isomorphic pereopods, larger posteri- 
orly, uncarinated. Pleon outline subtriangu- 
lar, slightly narrower than seventh pereonite, 
distinct pleonites separated laterally; four 
pairs of sessile ovoid pleopods conspicuous 
in ventral view; gross pleotelson triangular 
shape, wide basis and rounded tip (Fig. 10); 
uropods absent. 

Etymology.—The specific name refers to 
the Caribbean islands, where the examined 
specimens were collected. 

Variation in female specimens.—In 
younger specimens the cleft of the pleotel- 
son is open V-shaped, and the tips of the 
lobes reach to or beyond the tips of the fifth 
pleonite (Fig. 15). Size and shape of the 
carina of the pereopods varies according to 


its position. In P. insularis the shape of the 
carina is rectangular in PI—P2, convex in 
P3—P4, or subquadrangular in P5—P7, in- 
creasing its size behind. 

In adult specimens the palp of the max- 
illiped usually is prominent and finger-like, 
sometimes with a pair of distal setae; in oth- 
er specimens the shape of the maxilliped is 
subquadrate, rudimentary, reduced to a 
small, acute, lateral point, or bearing two 
digital processes directed outward. 

Variation in males.—TYhe body shape 
and size in males varies with the state of 
development (Figs. 12—14). The pleon in P. 
insularis is composed by four or five pleon- 
ites, depending upon the state of develop- 
ment, which are differentiated dorsally and 


VOLUME 114, NUMBER 4 


Table 1.—Extended. 


Characters 


Palp of maxilliped 
& number of setae 


Height & Shape 


of the carina Pleopods: length 


Digitiform; 2 Extremely high, | Exopodite large, 


setae subquadrangular § endopodite 
small 
Extremely high, Small 
quadrangular 
Pyramidal or Extremely high, Small 
ovoid; 4 setae rounded 
Medium, rounded 
Ovoid, 8-10 High, rounded Small 
setae 
High, rounded Small 
Medium, rounded Small or 
subequal 
Ovoid, 8-10 
setae 
Small 
High, Small 
quadrangular 
Ovoid, High, rounded Exopodite large, 
undetermined endopodite 
small 
Ovoid, 3 setae Small 


laterally, but sometimes the pleonites are 
differentiated only laterally. 

The shape of the pleotelson in males of 
P. insularis can be subtriangular, subovoid, 
subrounded, irregularly shaped, fused cen- 
tral and laterally with the previous pleonite 
on one or both sides but invariably bulbous 
and blunt, directed behind or laterally. 

Description of the epicaridium larva.— 
Body length: 0.18 to 0.25 mm; length width 
ratio from 1.7 to 2.3. Antenna large, six 
segmented, reaching between the second 
and sixth pleonites; segments 1—4 large; 
distal seta on internal face of fourth, fifth 
and sixth segments thinner than others. Dis- 
tal segment bordered by three small spines 
and two large unequal setae (Fig. 16). Max- 
illipeds not distinguishable. Pereopods sub- 


921 
Pleopods: protrusion 
from border of pleon Pleotelson Uropods 
Protruding far Fissured, lobes not Absent 
separated 
Protruding Entire, notched Absent 
slightly 
Diminished Notched Present 
Not protruding Fissured Rudimentary 
Protruding far Entire or slightly Absent 
notched 
Not protruding Slightly notched Absent 
Protrusion Entire or notched 
variable 
Notched 
Not protruding or Entire or notched 
only slightly 
Not protruding or Entire (right/ 
far rounded) or 
notched 
Diminished Notched Absent 
Protruding Bilobated 


slightly or not 


equal and dimorphic; first three pairs robust 
(Fig. 17); other three thinner (Fig. 18). Pos- 
teroventral border of fifth pleonite finely 
denticulate (Fig. 19); ventral side of sixth 
pleonite formed of plates. First pleopod 
with basal plate and a long, thin, curved 
external ramus ending in three long feath- 
ered setae. Posteromedial corner of basi- 
podite of pleopod 1 with large feathered se- 
tae (Fig. 20). This seta absent in following 
three pairs of pleopods, and replaced in fifth 
pair of pleopods by an ovoid branch twice 
the length of the exopodite. Fifth pair un- 
segmented, basipodite semi-rounded, exo- 
podite ending in three short feathered setae 
(Fig. 21). Uropods of two equal branches 
ending in two denticles and one strong me- 
dian seta (Fig. 22); anal tube small. 


9 


N 
N 


Macrobrachium, P. = Palaemonetes, Pa. = Palaemon). 


Table 2.—Morphological comparative characters for the American species of Probopyrus (Males). (Hosts: M. 


Characters 


Source 


Shape of pleon Shape of pleotelson 


Species/host 


Present work 


Subtriangular, digitiform, bulky 


Rounded, entire or fissured 


Scutelliform or subtriangular 


Semicircular or ovoid 


P. insularis/M. faustinum 


Richardson, 1904; 1905 


P. bithynis/M. ohionis (sic) 


P. bithynis/M. olfersii 


Roman-Contreras, 1996 


Richardson, 1912 


Rounded 


Subtrapezoidal 
Semicircular 


P. markhami/M. americanum 


P. meeki/M. acanthurus 


Button rounded 
Button rounded 


Rounded 


Roman-Contreras, 1993 


Richardson, 1912 


Semicircular 


P. pacificensis/M. tenellum 


Semicircular 


P. panamensis/M, acanthurus 
P. floridensis/P. paludosus 


P. pandalicola/P. hiltonii 
P. pandalicola/P. pugio 


Richardson, 1904; 1905 


Rounded or bare-necked 


Rounded 
Rounded 


Semicircular 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Jiménez and Vargas, 1990 


Markham, 1985 


Semicircular 
Variable 


Packard, 1879; Gissler, 1882; Bonnier, 1900; 


Rounded or triangular 


Semicircular, posterior edge 


P. pandalicola/P. vulgaris 


Richardson, 1905; Markham, 1985 


Verdi and Schuldt, 1988 


truncate 
Slightly rounded 


Rounded or truncate 


P. ringueleti/P. argentinus 


Lemos de Castro and Brasil Lima, 1974 


Rounded, fissured or bilobulated 


Wide, semicircular 


P. palaemoni/Pa. pandaliformis 


Discussion 


The use of host selectivity, pigmentation 
patterns, habitat, behavior, morphological 
structures, and statistical methods to differ- 
entiate species in the genus Probopyrus 
have been discussed in a recent paper by 
Roman-Contreras (1993, 1996). The fol- 
lowing morphological characters that 
should be described carefully in to differ- 
entiating species have been discussed by 
Giard & Bonnier (1888), Bonnier (1900), 
Richardson (1904, 1905, 1912), Chopra 
(1923), and Roman-Contreras (1993, 1996). 

Females of P. insularis can be differen- 
tiated from other American species of Pro- 
bopyrus by their relatively short and narrow 
pleotelson. In adult specimens a deep fis- 
sure with rounded and unseparated lobes is 
present; usually the edge of the lobes do not 
reach the tip of the fifth pleonite. The shape 
of the female pleotelson of P. insularis is 
similar only to P. meeki Richardson, 1912. 
However, the sixth segment of P. meeki has 
a posterior notch that extends half of its 
length and forming two posterior unsepa- 
rated lobes (Richardson, 1912: fig. 1). 

The robust carina of the seventh leg of 
P. insularis is comparable to the high and 
prominent carina of P. markhami. All legs 
of P. bithynis Richardson, 1904 have an ex- 
tremely high carina (Richardson 1904, 
1905). The legs of P. pandalicola have a 
high quadrangular-shaped carina, while the 
carina of P. floridensis Richardson, 1904 
are rounded (Richardson 1912). Romdan- 
Contreras (1993, 1996) noted the rounded 
carina of P. pacificensis. 

Although the maxilliped palp of Probo- 
pyrus may vary according to the develop- 
mental state of the specimens, the mor- 
phology has proven valuable in separating 
species. The maxilliped palp of adult fe- 
males of P. insularis is finger-like in most 
specimens (79%); a small percentage (14%) 
have two setae on the tip or no palp is pre- 
sent; few individuals have a triangular, a 
rounded, or a rudimentary palp. For Pro- 
bopyrus ringueleti Verdi & Schuldt, 1988, 


VOLUME 114, NUMBER 4 923 


GE ab 


Figs. 1-5. Holotype female of Probopyrus insularis sp. nov. 1), in ventral view (oostegite 1 and barbula 
removed); 2), same in dorsal view; 3), antennula and antenna from a paratype specimen (picture taken with 
scanning electron microscopy: SEM); 4), maxilliped in dorsal view; 5), barbula of a paratype. Scale bar: 1 and 
2 = 2.0 mm; 4 = 0.5 mm; 5 = 1.0 mm. 


924 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 6-8. Probopyrus insularis sp. nov. 6), oostegite 1 in dorsal view; 7), pereopod 7 of holotype female; 
8), pereopods 1—7 of paratype female. Scale bar: 6 and 15 = 1.0 mm; 7 = 0.5 mm. 


VOLUME 114, NUMBER 4 


Ne) 


25 


Figs. 9-15. Probopyrus insularis sp. noy. 9), pleopods 1—5 of paratype female; 10), allotype male in ventral 
view; 11), antennula and antenna of paratype male (taken with SEM); 12-14), variation of males in dorsal view; 
15), young female from Puerto Rico (USNM 235988). Scale bar: 8 and 10 = 0.5 mm; 11 = 0.05 mm; 12-14 
= 1.0 mm. 


926 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


17 


ao Be 


18 49 


Figs. 16-22. Epicardium larva of Probopyrus insularis sp. nov. 16), antenna; 17), pereopod 1; 18), pe- 
reopod 4; 19), posterior edge of the fifth pleonite; 20), first pleopod; 21), fifth pleopod; 22), uropods. Scale 
bar: 0.05 mm. 


VOLUME 114, NUMBER 4 


the palp is rectangular and bears an unde- 
termined number of setae. The palp of P. 
pacificensis is ovoid bearing 8 to 10 setae 
(Romén-Contreras 1993), while the palp of 
P. markhami is pyramidal and bears a re- 
duced number of setae (Roman-Contreras 
1996). 

The pleon of males also has been used to 
differentiate some species of Probopyrus. 
However, the narrow pleon in males of P. 
insularis is not unique; it is comparable to 
that of P. meeki, and the gross structure of 
the pleotelson is similar to adult specimens 
of P. meeki as well as P. markhami. 

In addition to structures of adult females 
and males, larval morphology of Probopy- 
rus has been used to separate species (Dale 
& Anderson 1982). Although such features 
may prove to be important, mature larval 
stages are not always available to taxono- 
mists. 


Acknowledgments 


Special thanks are given to Y. Thérézien 
and E. Fiévet for collecting the specimens 
from Guadeloupe, and making them avail- 
able to us; to B. Kensley, J. Clark, and M. 
Schotte (NMNH, Smithsonian Institution, 
Washington, D.C.) for the loan of cata- 
logued specimens and facilities during sev- 
eral short visits of RRC to the Division; to 
M. Martinez-Mayén (ICMyL-UNAM) for 
his technical assistance in the laboratory; to 
J. C. Markham, and three anonymous ref- 
erees who helped improve the final manu- 
script; and to A. E. Viniegra for the art 
work. 


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Pereira S., G. A. 1991. Camarones de agua dulce de 
Venezuela II: Nuevas adiciones en las familias 
Atyidae y Palaemonidae (Crustacea, Decapoda, 
Caridea).—Acta Biolégica Venezuélica 13:75— 
88. 

Richardson, H. 1904. Contributions to the natural his- 
tory of the Isopoda.—Proceedings of the United 
States National Museum 27:58—89. 

. 1905. Monograph on the isopods of North 

America.—Bulletin of the United States Nation- 

al Museum 54:1—727. 

. 1912. Descriptions of two new parasitic iso- 
pods belonging to the genera Palaegyge and 
Probopyrus from Panama.—Proceedings of the 
United States National Museum 42:521—524. 

Roman-Contreras, R. 1993. Probopyrus pacificensis, a 
new parasite species (Isopoda: Bopyridae) of 
Macrobrachium tenellum (Smith, 1871) (Deca- 
poda: Palaemonidae) of the Pacific coast of 
México.—Proceedings of the Biological Socie- 
ty of Washington 106:689—697. 

. 1996. A new species of Probopyrus (Isopoda, 
Bopyridae), parasite of Macrobrachium ameri- 
canum Bate, 1868 (Decapoda, Palaemoni- 
dae).—Crustaceana 69:204—210. 

Saussure, H. de. 1857. Diagnoses de quelques Crus- 
tacés noveaux de |’ Amérique tropicale.—Revue 
et Magasine de Zoologie 2:501—505. 

Verdi, A. C., & M. Schuldt. 1988. Descripcion y biol- 
ogia de Probopyrus ringueleti n. sp. (Crustacea, 
Epicaridea, Bopyridae). Efectos sobre el hos- 
pedador Palaemonetes argentinus Nobili (Crus- 
tacea, Caridea, Palaemonidae).—Anales de la 
Sociedad Cientifica Argentina 218:15—26. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(4):929—937. 2001. 


A new asellote isopod of the genus Santia Sivertsen & Holthuis, 1980 
(Crustacea: Isopoda: Aseliota: Santiidae) from Japan 


Michitaka Shimomura and Shunsuke KF Mawatari 


Division of Biological Sciences, Graduate School of Science, Hokkaido University, North 10, 
West 8, Kita-ku, Sapporo 060-0810, Japan 


Abstract.—Santia longisetosa, a new species of the family Santiidae (Iso- 
poda: Asellota) is described from the Pacific coast of Shikoku, Japan. The new 
species differs from its congeners in having a long robust sensory seta distal- 
ventrally on each propodus of pereopods 2-7, the nearly straight frontal margin 
of the head bearing some long setae, and 2 long robust sensory setae distal- 


laterally on article 2 of antenna 1. 


Santiidae is a small family of four genera 
and 21 species in the suborder Asellota. 
Santia Sivertsen & Holthuis, 1980, is the 
largest genus in the family and includes 15 
species (Wolff 1989, Wolff & Brandt 2000, 
Shimomura & Mawatari 2000), all from 
marine benthic habitats. Among them only 
one species, Santia katoi Shimomura & 
Mawatari, 2000 has been so far described 
from Japan. Our recent investigation yield- 
ed an undescribed species of Santia from 
the subtidal zone of Kochi Prefecture, Shi- 
koku, the second record of the genus from 
Japan. 


Materials and Methods 


Specimens were obtained from 0.5 m 
depth of the subtidal zone by filtering sur- 
face water; specimens were fixed with 5% 
neutral formalin solution diluted with sea- 
water, and preserved in 70% ethanol. Each 
isopod was dissected and prepared for ob- 
servation using a light microscope equipped 
with Nomarski differential interference 
contrast (Shimomura & Mawatari 1999). 
Total length as indicated in “‘Material ex- 
amined”’’ was measured from the tip of the 
head to the end of the pleotelson. 

The. type specimens are deposited in the 
Zoological Museum, Division of Biological 
Sciences, Graduate School of Science, Hok- 


kaido University (ZIHU), and in the Toya- 
ma Science Museum (TOYA). 


Santia longisetosa, new species 
Figs. 1-4 


Material examined.—Murotomisaki-cho, 
Muroto Cape, Kochi Prefecture, Japan, 
33°16'N, 134°7’E, 0.5 m, surface water, 
subtidal, 27 June 2000: holotype, male, 1.0 
mm (ZIHU-01963); paratype, ovig. female, 
1.1 mm (TOYA Cr-12877); paratypes, 2 fe- 
males, 1.1 mm (TOYA Cr-12878), 1.3 mm 
(ZIHU-01966). 

Description.—Male: Body (Fig. 1A) 
about 2.4 times as long as maximum width. 
Head 2.0 times as broad as long, slightly 
narrower than pereonite |, with 10 dorsal 
setae; frontal margin of head nearly 
straight, with 8 long setae; labrum surpass- 
ing head anteriorly; posterior margin of 
head convex between eyestalks. Preocular 
lobes narrow, with | or 2 apical setae. Eyes 
dorsal lateral, each with 3 ommatidia. Per- 
eonites 1—4 laterally rounded, each with 4— 
7 lateral and few dorsal setae; pereonites 5— 
7 laterally rounded, each with O-—3 lateral 
and 2 dorsal setae. Pereonites 1, 2 and 4 
subequal in length; pereonite 3 slightly 
shorter than pereonite 2; pereonites 5 to 7 
increasing in length. Pereonites 1 and 2 
subequal in width; pereonite 3 slightly wid- 


930 


er than pereonite 2; pereonite 4 as broad as 
pereonite 3; pereonite 5 widest; pereonites 
5 to 7 decreasing in width. Coxal plates 
dorsally visible on pereonites 4—7, laterally 
rounded, each with 2 or 3 lateral setae. 
Pleonite short and narrow, without dorsal 
setae. Pleotelson (Figs. 1A, 2A) about 1.1 
times as long as broad, with 13 dorsal, 2 
posterolateral, 2 apical, and 12 ventral se- 
tae. Uropod (Fig. 2B) stout, directed pos- 
teriorly, shorter than pleotelson. Protopod 
wide posteriorly, about *% as long as exo- 
pod, with 1 mesial and | lateral robust sen- 
sory setae. Exopod and endopod longer 
than broad. Exopod about 1.5 times as long 
as protopod, with 1 ventral, 5 dorsal, 2 lat- 
eral, | subapical and 4 apical robust sensory 
setae, with 1 apical simple seta; endopod as 
long as peduncle, with 2 ventral, 1 dorsal, 
2 subapical and 2 apical robust sensory se- 
tae, with 3 lateral and 2 subapical plumose 
setae. 

Antenna 1 (Fig. 2C) of 6 articles. Article 
1 broadest, with 1 distal-lateral and 1 distal- 
medial setae; article 2 longest, narrower 
than article 1, with | distal-ventral, 1 distal- 
dorsal and 1 lateral simple setae, with 1 dis- 
tal-lateral and 1 distal-medial plumose se- 
tae, and distal-lateral with 2 long robust 
sensory setae; article 3 narrow, with 1 dis- 
tal-ventral and 1 distal-dorsal setae; article 
4 slightly shorter than article 3, without se- 
tae; article 5 about 1.7 times as long as ar- 
ticle 4, apically with 1 aesthetasc; article 6 
narrow and short, apically bearing | aesth- 
etasc and 2 short, 1 long setae. 

Antenna 2 (Fig. 2E) composed of 6 stout 
articles and 12 thin flagellar articles. Article 
1 with 1 distal-lateral seta; article 2 as broad 
as article 1, without setae; article 3 longer 
than article 2, with 3 distal-ventral, 1 dorsal 
and 1 distal-medial setae; article 4 shorter 
than article 3, with 2 distal-ventral, 3 distal- 
lateral and 1 medial setae; article 5 as long 
as article 6, with 2 ventral, 3 dorsal, 2 lat- 
eral and 3 mesial simple setae, and with | 
long robust sensory seta distal-medially and 
1 plumose seta laterally; article 6 narrower 
than article 5, bearing 3 ventral, 4 dorsal, 2 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


lateral and 2 medial simple setae, and with 
1 lateral and 1 medial plumose setae. Fla- 
gellum about 2.6 times as long as article 6, 
each with many setae. 

Left mandible (Fig. 2F) lacking palp, 
bearing | short seta arising from its base; 
incisor with 4 cusps; lacinia mobilis with 3 
teeth; setal row with 4 setae; molar process 
stout, bearing 2 apical setae. Right mandi- 
ble (Fig. 2G) lacking palp and lacinia mob- 
ilis, bearing | short seta arising from its 
base; incisor with 6 cusps; setal row with 4 
setae; molar process stout, bearing 2 apical 
setae. 

Maxilla | (Fig. 2H) with inner lobe bear- 
ing 4 apical setae; outer lobe apically with 
5 pectinate setae and 5 simple setae, dor- 
sally with many short setae. Maxilla 2 (Fig. 
21) with inner lobe with 5 apical setae and 
many dorsal setae; outer 2 lobes each with 
4 apical setae. 

Maxilliped (Fig. 3A) with moderately 
broad palp composed of 5 articles; article 1 
as broad as article 2, with | medial seta; 
article 2 trapezoidal, about 2.3 times as long 
as article 1, with | lateral and 2 medial se- 
tae; article 3 as long as article 2, with 2 
lateral and 3 medial setae; article 4 slightly 
shorter than article 3, with 2 lateral and 4 
medial setae; article 5 narrowest, with 1 
subapical and 2 apical setae; endite quad- 
rate, bearing 4 dorsal, 6 distal simple setae, 
with 5 distal pectinate setae and 3 subdistal 
fan-shaped setae, and with many short setae 
laterally and 2 coupling hooks medially, 
epipod lanceolate, moderately broad, nar- 
rower than endite, with rounded apex. 

Pereopod | (Fig. 3B) shorter than pereo- 
pods 2-7: basis the longest article, with 2 
ventral, 2 dorsal and 1 lateral setae; ischium 
narrower than basis, bearing | ventral, 2 
dorsal, 1 lateral and 2 distal setae; merus 
trapezoidal, with | ventral, 3 distal-ventral 
and 2 distal-dorsal simple setae, with 1 long 
robust sensory seta distal-dorsally; carpus 
trapezoidal, broadest, ventrally with 2 long 
robust sensory setae and 3 simple setae, 
medially with | simple seta and 3 spinulose 
scales, and with 1 simple seta distal-dorsal- 


VOLUME 114, NUMBER 4 


931 


Fig. 1. 
dorsal. Scales = 0.1 mm. 


ly; propodus ovate, ventrally with 5 simple 
setae and | robust sensory setae, medially 
with 1 simple short seta and 7 spinulose 
scales, and with 5 simple setae dorsally; 
dactylus shorter than propodus, narrowest 
of all articles, with 2 distolateral and 3 me- 
dially setae, 1 curved unguis, and 1 short 
accessory spine. 

Pereopod 2 (Fig. 3C) as long as pereopod 


Santia longisetosa, new species. A, holotype male, dorsal; B, paratype female (TOYA Cr-12877), 


4: basis with 2 ventral, 2 dorsal and 1 lat- 
eral setae; ischium shorter than basis, with 
2 ventral, 2 dorsal and 2 medially setae; 
merus trapezoidal, distal-dorsally with 2 
simple and | long robust sensory setae, 
with 2 ventral, 1 distal-lateral and 1 distal- 
medial simple setae; carpus longer than ba- 
sis, dorsally with 4 simple, 1 robust sensory 
and | plumose setae, with 3 ventral, 2 lat- 


932 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Santia longisetosa, new species. A-C, E-I, holotype male; D, paratype female (ZIHU-01966): A, 
pereonite 7 and pleon, ventral; B, right uropod, ventral; C, left antenna 1, dorsal; D, left antenna 1, dorsal; E, 
left antenna 2, dorsal; E left mandible, dorsal; G, right mandible, dorsal; H, right maxilla 1, ventral; left maxilla 
2, ventral. Scales = 0.1 mm. 


VOLUME 114, NUMBER 4 933 


Fig. 3. Santia longisetosa, new species. A-G, holotype male: A, left maxilliped, dorsal; B, left pereopod 1, 
mesial; C, right pereopod 2, mesial; D, right pereopod 3, mesial; E, left pereopod 4, lateral; E left pereopod 5, 
lateral; G, right pereopod 6, mesial. Scales = 0.1 mm. 


934 


eral and 2 medial simple setae; propodus 
the longest article, dorsally with 5 simple 
and | plumose setae, with 3 ventral, 3 lat- 
eral, 4 mesial simple setae, and with 1 dis- 
tal-ventrally long robust sensory seta and 
many ventral short setae; dactylus the nar- 
rowest article; with 2 distal-lateral and 3 
medial setae, 1 curved unguis, and | minute 
accessory spine. 

Pereopod 3 (Fig. 3D) slightly longer than 
pereopod 2: basis with 2 ventral and 2 dor- 
sal setae; ischium with 2 ventral, 2 dorsal 
and | distal-medial setae; merus distal-dor- 
sally with 1 simple and 1 long robust sen- 
sory setae, with 2 ventral, 1 distal-medial 
and | distal-lateral simple setae; carpus dor- 
sally with 4 simple, | robust sensory and 1 
plumose setae, with 4 ventral, 2 lateral and 
2 mesial simple setae; propodus dorsally 
with 10 simple and 1 plumose setae, with 
2 ventral, 1 lateral, 1 mesial simple setae, 
and with 1 ventral-distal long robust sen- 
sory, 1 ventral short robust sensory setae 
and many ventral short setae; dactylus with 
2 distal-lateral and 3 medial setae, 1 curved 
unguis, and | minute accessory spine. 

Pereopod 4 (Fig. 3E) slightly shorter than 
pereopod 3: basis with 1 ventral and 2 dor- 
sal setae; ischium with 2 ventral, 2 dorsal 
and 1 distal-medial setae; merus distal-dor- 
sal with 1 simple and | long robust sensory 
setae, with | distal-ventral and 1 distal-lat- 
eral simple setae; carpus dorsally with 1 
simple, 1 long robust sensory and 1 plu- 
mose setae, with 2 ventral, 3 lateral and | 
mesial simple setae; propodus dorsally with 
4 simple and 1 plumose setae, with 2 ven- 
tral and | lateral simple setae, and with 1 
distal-ventral long robust sensory seta and 
many ventral short setae; dactylus with 2 
distal-lateral and 3 medial setae, | curved 
unguis, and 1 minute accessory spine. 

Pereopod 5 (Fig. 3F) longer than pereo- 
pod 3: basis dorsally with 2 simple and 1 
plumose setae, with 3 ventral simple setae; 
ischium with 1 dorsal robust sensory seta 
and 3 ventral, 2 lateral and | distal-medial 
simple setae; merus distal-dorsal with 1 
simple and 1 long robust sensory setae, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


with 2 ventral, 1 distal-medial and 1 distal- 
lateral simple setae; carpus dorsally with 2 
simple, 2 long robust sensory, 1 short ro- 
bust sensory and 1 plumose setae, with 1 
ventral robust sensory seta and 2 ventral, 2 
lateral and 2 medial simple setae; propodus 
dorsally with 7 simple and | plumose setae, 
with 3 ventral, 3 lateral, 2 medial simple 
setae, and with | distal-ventral long robust 
sensory, | ventral short robust sensory setae 
and many ventral short setae; dactylus with 
2 distal-lateral and 3 medial setae, 1 curved 
unguis, and | minute accessory spine. 
Pereopod 6 (Fig. 3F) the longest pereo- 
pod: basis dorsally with 2 simple and 1 plu- 
mose setae, with 3 ventral simple setae; is- 
chium with 1| dorsal robust sensory seta and 
3 ventral, 3 medial simple setae; merus dis- 
tal-dorsal with 1 simple and 1 long robust 
sensory setae, with 2 ventral and 1 distal- 
medial simple setae; carpus dorsally with 1 
simple, 2 long robust sensory, 2 short ro- 
bust sensory setae and 1 plumose setae, 
with 2 ventral robust sensory setae and 2 
ventral, 2 medial simple setae; propodus 
dorsally with 7 simple and | plumose setae, 
with 1 distal-ventral and 1 lateral simple se- 
tae, and with 1 distal-ventral long robust 
sensory, 2 ventral short robust sensory setae 
and many ventral short setae; dactylus with 
2 distal-lateral and 3 medial setae, 1 curved 
unguis, and 1 minute accessory spine. 
Pereopod 7 (Fig. 4A) slightly shorter 
than pereopod 6: basis dorsally with 1 sim- 
ple and 2 plumose setae, with 2 ventral sim- 
ple setae; ischium with 1 dorsal robust sen- 
sory seta and 2 ventral, 2 dorsal and 1 dis- 
tal-lateral simple setae; merus distal-dorsal 
with 1 simple and 1 long robust sensory 
setae, with 1 distal-ventral, 1 distal-medial 
and | distal-lateral simple setae; carpus dor- 
sally with 1 simple, 1 long robust sensory, 
2 short robust sensory setae and 1 plumose 
setae, with | short ventral robust sensory, | 
distal-lateral short robust sensory setae, and 
with 2 ventral, | lateral and 2 medial simple 
setae; propodus, dorsally with 7 simple and 
1 plumose setae, with 1 distal-ventral, 2 lat- 
eral simple setae, and with | distal-ventral 


VOLUME 114, NUMBER 4 


935, 


Fig. 4. Santia longisetosa, new species. A-E, holotype male; EK paratype female (ZIHU-01966): A, right 
pereopod 7, lateral; B, left pereopod 2, ventral; C, left pleopod 3, ventral; D, right pleopod 4, dorsal; E, left 
pleopod 5, dorsal; K operculum, ventral. Scales = 0.1 mm. 


long robust sensory, 3 distal-ventral short 
robust sensory setae, many ventral short se- 
tae and 7 lateral spinulose scales; dactylus 
with 2 distal-lateral and 3 medial setae, 1 
curved unguis, and 1 minute accessory 
spine. 

Pleopod 1 broken. Pleopod 2 (Fig. 4B) 
with broad protopod, tapering to rounded 
apex, lateral margin convex, bearing | api- 
cal and 3 submarginal setae; endopod with 
slender second article; exopod narrow. Ple- 
opod 3 (Fig. 4C) with endopod bearing 3 
stout, plumose setae distally; exopod com- 
posed of 2 articles, narrower than endopod; 
article 2 bearing 2 lateral simple setae. Ple- 
opod 4 (Fig. 4D) with exopod narrow, dis- 
tally with 1 stout, long plumose seta, lat- 


erally with many short setae; endopod ovate 
and broad, without setae. Pleopod 5 (Fig. 
4E) ovate, uniramous, about 1.6 times as 
long as broad, without setae. 

Female: Similar to male in morphology 
of all pereonal appendages. Body (Fig. 1B) 
about 2.1 times as long as maximum width. 
Head slightly narrower than pereonite 1, 
with 10 dorsal setae; frontal margin of head 
nearly straight, with 9 long setae. Pereoni- 
tes 1—4 laterally rounded, each with 6-8 lat- 
eral setae and few dorsal setae; pereonites 
5-7 laterally rounded, each with 1-3 lateral 
setae and few dorsal setae. Pereonite | 
shorter than pereonite 2; pereonites 2 and 3 
subequal in length; pereonite 4 longest; per- 
eonites 5 and 6 subequal in length; pereon- 


936 


ite 7 longer than pereonite 6. Pereonites | 
to 3 increasing in width; pereonite 3 widest; 
pereonite 4 slightly narrower than pereonite 
3; pereonites 5 to 7 decreasing in width. 
Coxal plates dorsally visible on pereonites 
4—7, each with 1—3 lateral setae. Pleonite 
short and narrow, without dorsal setae. An- 
tenna | (Fig. 2D) composed of 5 articles. 
Article 1 broadest, with | distal-lateral and 
1 distal-medial setae; article 2 longest, nar- 
rower than article 1, with 1 distal-ventral, | 
lateral and 1 distal-medial simple setae, 
with 1 distal-lateral plumose seta, and with 
2 distal-lateral robust sensory setae; article 
3 narrow, with | distal-ventral seta; article 
4 slightly shorter than article 3, without se- 
tae; article 5 about 1.6 times as long as ar- 
ticle 4, subapically with | short setae, api- 
cally with | aesthetasc and 2 short, | long 
setae. Operculum (Fig. 4F) about 1.2 times 
as long as broad, apically with short round- 
ed projection, submarginally with 14 setae. 
Etymology.—The specific name refers to 
the long sensory setae on the antenna | and 
on the propodi of the pereopods 2-7. 
Remarks.—The present new species is 
assigned to Santiidae Kussakin, 1988, hav- 
ing a set of the following characters: eyes 
situated on lateral processes, short antenna 
1 having 5 or 6 articles, antenna 2 with 4 
short proximal, 2 long distal articles and 
flagellum, subcylindrical truncate mandib- 
ular molar process, narrow maxillipedal 
palp, distally pointed epipod of maxilliped, 
coxal plates of pereonites 5—7 visible dor- 
sally, prehensile pereopod 1 and ambula- 
tory pereopods 2—7, uropods situated on 
posterolateral margin of pleotelson, and ter- 
minally exposed anus. The following fea- 
tures displayed by the present new species 
indicate that it belongs to Santia Sivertsen 
& Holthuis, 1980: antenna 1 and 2 in an- 
terior indentations laterally on the head, 
preocular lobes in front of the eyestalks, 
pleotelson apically with short rounded pro- 
jection, pereopod | armed with 1 unguis 
and | accessory spine, female operculum 
longer than broad, and uropods with stout 
protopod, cylindrical endopod and exopod. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Santia longisetosa is distinguishable 
from its congeners by 2 long robust sensory 
setae on article 2 of antenna 1 and 1| long 
robust sensory seta ventrodistally on each 
propodus of pereopods 2—7. The nearly © 
straight frontal margin of head and the ro- 
bust long uropods are shared by Santia lon- 
gisetosa, S. milleri (Menzies & Glynn, 
1968) from Caribbean Sea (type locality), 
S. hispida (Vanhoffen, 1914) from St. Paul 
Island, the southern Indian Ocean (type lo- 
cality), Tristan da Cunha (Nordenstam 
1933) and the Magellan Strait (Winkler 
1993). The present new species is distin- 
guished from S. milleri by the following 
features (those of S. milleri in parentheses): 
article 2 of antenna | has 2 long robust sen- 
sory setae, propodi of pereopods 2—7 have 
a long robust sensory seta distal-ventral, 
pereonite | is slightly wider than head (nar- 
rower than head), coxal plates lack anterior- 
lateral stout setae (each with anterior-lateral 
stout seta), mandibular palp is absent (pre- 
sent), maxillipedal palp is moderately broad 
(narrow). Santia hispida differs from the 
present new species in having the narrow 
maxillipedal palp, the mandibular palp, 
many long stout dorsal and lateral setae on 
head, pereon and pleon, very long second 
article of pleopod | in male, and a pair of 
stout setae on operculum in female. 

Santia charcoti (Richardson, 1906) 
(Hodgson 1910, Wilson 1980) from the 
Antarctic has stout uropods, similar to those 
of S. longisetosa. The present new species 
is however distinguished from the S. char- 
coti by the following features (those of S. 
longisetosa in parentheses): the frontal mar- 
gin of head is bilobed (unilobed), many 
long setae are on head, pereon and pleon 
(some short setae), the pleotelson is short 
(long), endopod of uropod is curved 
(straight), protopod of pleopod 1 in male is 
broad (narrow), and coxal plates of pereon- 
ite 7 are dorsally invisible (visible). 

The present new species differs from an- 
other Japanese species, S. katoi Shimomura 
& Mawatari, 2000 described from Shira- 
hama coast, Wakayama Prefecture in the 


VOLUME 114, NUMBER 4 


following features (those of S. katoi in pa- 
rentheses): the mandible lacks palp (palp 
present), the frontal margin of head is near- 
ly straight (slightly convex), with 8 or 9 
long setae (4 short setae), the head is nar- 
rower than pereopod 1 (broader than pereo- 
pod 1), the uropod shorter than pleotelson 
(onger than pleotelson), the article 2 of an- 
tenna | has 2 long robust sensory setae dis- 
tolaterally (without sensory setae), and the 
propodi of pereopods 2—7 are armed with 
long robust sensory setae ventrodistally 
(with short robust sensory setae). 

The present new species shows a varia- 
tion in segmentation of antenna | as fol- 
lows: holotype male has 6-articulate anten- 
na 1 consisting of 2 stout, 2 short, 1 long 
and | minute articles, while all paratype fe- 
males have 5-articulate antenna | consisting 
of 2 stout, 2 short and 1 long articles. 


Acknowledgments 


We thank two anonymous reviewers for 
their critical reading of the manuscript. 


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strupia 15(7):177-191. 

, & A, Brandt. 2000. Caribbean species of 

Munnidae, Paramunnidae and Santiidae (Iso- 

poda: Asellota).—Steenstrupia 25(1):121—146. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(4):938—943. 2001. 


A new genus and species of freshwater crab from Colombia 
(Crustacea: Decapoda: Pseudothelphusidae) 


Martha R. Campos 


Universidad Nacional de Colombia, Instituto de Ciencias Naturales, Apartado Aéreo 103698, 
Bogota, D.C., Colombia, S. A., e-mail: dcamposr@cable.net.co 


. Abstract.—A new genus, Achagua, is established for the new species A. 
casanarensis, and Eudaniela pestai (Pretzmann). Consequently, the latter is 
removed from the genus Eudaniela Pretzmann. Species of the new genus are 
known exclusively from the upper Colombian Orinoco basin and the western 
region of the Venezuelan coastal cordillera. The new genus is characterized by 
features of the third maxilliped, the orifice of branchial efferent channel, and 


the first male gonopod. 


During faunistic surveys of the upper 
Colombian Orinoco basin, specimens be- 
longing to a new species of the family Pse- 
dothelphusidae were collected. The new 
species resembles Eudaniela pestai (Pretz- 
mann, 1965) which according to Rodriguez 
(1982) and Rodriguez & Pereira (1992), ex- 
hibits the least derived first male gonopod 
morphology and less reduced exognath of 
the third maxilliped within the tribe King- 
sleyini. These two species are sufficiently 
distinct from others of the tribe in the char- 
acteristics of the third maxilliped, orifice of 
the branchial efferent channel, and first 
male gonopod, to warrant their assignment 
to a new genus. Eudaniela pestai is distrib- 
uted in the western region of the Venezue- 
lan coastal cordillera. The new genus and 
species are described herein. The new ge- 
nus is considered a basal group from which 
species of Eudaniela Pretzmann, 1971 orig- 
inated (Rodriguez, pers. comm.). 

The terminology used for the morpholo- 
gy of the male first gonopod follows Smal- 
ley (1964) and Rodriguez (1982). The ab- 
breviations cl and cb stand for carapace 
length and carapace breadth, respectively. 
The color nomenclature used follows Smi- 
the (1975). The material is deposited in 
Museo de Historia Natural, Instituto de 


Ciencias Naturales, Universidad Nacional 
de Colombia, Bogota (ICN-MHN). 


Systematics 


Family Pseudothelphusidae Rathbun, 1893 
Tribe Kingsleyini Bott, 1970 
Genus Achagua, new genus 


Diagnosis.—Third maxilliped with mer- 
us of endognath regularly curved; exognath 
approximately 0.5 times length of ischium; 
orifice of branchial efferent channel partial- 
ly closed by spine of jugal angle, and by 
production of lateral lobe of epistome. First 
male gonopod straight with wide marginal 
process; subapical mesial process spine- 
like; mesial plate quadrate or slightly 
rounded, apex with mesial and cephalic 
plates distally overlapping or in parallel ar- 
rangement. 

Type species.—Achagua casanarensis, 
new species. 

Other included species.—Eudaniela pes- 
tai (Pretzmann, 1965) 

Etymology.—tThe genus is named in hon- 
or of the Achagua Indians, who lived in the 
region where the new genus was discov- 
ered. Gender: feminine. 


VOLUME 114, NUMBER 4 


Achagua casanarensis, new species 
Bigs. Ie 2 


Holotype.—Municipio Aguazul, 26 km 
SW from Yopal, Casanare Department, Co- 
lombia, 290 m alt., 3 Oct 1995, leg E Fer- 
nandez: | male, cl 32.7 mm, cb 51.6 mm, 
ICN-MHN-CR 1626. 

Paratype.—Same locality data as holo- 
type: 1 male, cl 29.3 mm, cb 48.7 mm, 
ICN-MHN-CR 1862. 

Type locality.—Municipio Aguazul, 26 
km SW from Yopal, Casanare Department, 
Colombia. 

Diagnosis.—Marginal process of first 
male gonopod wide, spatulated distally; 
with subapical broad-base, and distally 
acute spine-like mesial process; mesial 
plate quadrate; apex with mesial and ce- 
phalic plates distally overlapping, and basal 
triangular projection; field of spines narrow 
and straight, spinules conspicuous, uniform 
in size, directed distally. 

Description of holotype.—Carapace (Fig. 
1A) with deep, straight cervical groove, 
ending short distance from lateral margin. 
Anterolateral margin with depression be- 
hind external orbital angle, followed by 15 
tubercles on anterior half; posterior lateral 
margin smooth. Postfrontal lobes small, 
rounded, without anterior depressions; me- 
dian groove narrow, shallow, with incision 
on upper margin of front. Surface of cara- 
pace in front of postfrontal lobes flat and 
inclined anteriorly. Upper border of front 
crest-like, marked with row of tubercles; 
lower margin slightly sinuous in frontal 
view. Surface of front between upper and 
lower borders high and slightly excavated. 
Upper and lower orbital margins each with 
row of tubercles. Surface of carapace cov- 
ered with small papillae; limits between re- 
gions demarcated. Third maxilliped (Fig. 
2F) with merus of endognath regularly 
curved; exognath approximately 0.5 times 
length of ischium of third maxilliped. Ori- 
fice of branchial efferent channel partially 
closed by spine of jugal angle, and by pro- 
jection of lateral lobe of epistome (Fig. 1C). 


939 


First pereiopods heterochelous, right che- 
liped larger than left. Merus with 3 crests 
as follows: upper crest with rows of tuber- 
cles, internal lower crest with rows of teeth, 
and external lower crest with few tubercles. 
Carpus with 3 tubercles on internal crest, 
and prominent acute spine distally. Palm of 
larger cheliped swollen, without external 
tubercle, fingers gaping when closed, with 
rows of tubercles on dorsal side (Fig. 1B); 
smaller cheliped slightly swollen, fingers 
not gaping when closed. 

Walking legs (pereiopods 2—5) thick 
(Fig. 1A). Dactyli elongated, each about 1.6 
times as long as propodi, with papillae and 
5 longitudinal rows of large, dark spines di- 
minishing in size proximally. Number of 
spines and papillae on each dactylus ar- 
ranged as follows: | anterolateral row and 
1 anteroventral row each with 6 spines; 1 
external row with 6 spines and 1 pair of 
proximal papillae; 1 posterolateral row and 
1 posteroventral row with 4 spines. 

First male gonopod (Fig. 2A—E) straight, 
marginal process wide, spatulated distally; 
mesial border convex with subapical broad- 
based, and acute spine-like mesial process, 
followed by deep depression, ending distal- 
ly in quadrate mesial plate; lateral border 
slightly sinuous with notch at basis of sper- 
matic channel. Apex laterally, with mesial 
and cephalic plates distally overlapping, 
and basal triangular projection; field of 
spines narrow and straight, spinules con- 
spicuous, uniform in size, and directed dis- 
tally (Fig. 2B, D, E). 

Color.—In alcohol, the dorsal side of the 
carapace is gray (near 45, Smoke Gray) 
with Buff (24) specks. The walking legs are 
Tawny (38). The chelae are Pale Horn Col- 
or (92). The ventral surface is gray (near 
45, Smoke Gray) with Cinnamon (39) 
specks. 

Etymology.—The specific name refers to 
Casanare Department, where the specimens 
were collected. 

Remarks.—tThe new species is most sim- 
ilar to Achagua pestai. The two can be dis- 
tinguished by differences in the first male 


940 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


lcm UCPinzoy 


mm 


Fig. 1. Achagua casanarensis, new species, male holotype, cl 32.7 mm, cb 51.6 mm, ICN-MHN-CR 1626. 
A, dorsal view; B, chela of large cheliped, external view; C, orifice of branchial efferent channel. 


VOLUME 114, NUMBER 4 941 


SRivesy 
cm Lmm! 
Fig. 2. Achagua casanarensis, new species, male holotype, cl 32.7 mm, cb 51.6 mm, ICN-MHN-CR 1626. 


A, left first gonopod, caudal view; B, same, lateral view; C, same, cephalic view; D, same mesial view; E, apex 
of same, distal view; EF left third maxilliped, external view. 


942 


gonopod. In the new species the mesial 
plate is quadrate, while in A. pestai the me- 
sial plate is slightly rounded; the subapical 
mesial process is acute distally, followed by 
a deep depression in A. casanarensis, 
whereas the subapical mesial process is 
rounded, and there is a shallow depression 
in A. pestai. The marginal process is spat- 
ulated distally in the new species, whereas 
in A. pestai the marginal process is rounded 
distally. In the new species the apex has the 
mesial and cephalic plates distally overlap- 
ping, whereas they are parallel in A. pestai; 
in the new species there is a basal triangular 
projection in the apex which is absent in A. 
pestai. In A. casanarensis the field of spines 
is straight, and there are only uniform con- 
spicuous spines, whereas in A. pestai the 
field of spines is recurved, with large setae 
distally, and short spinules near the sper- 
matic channel. 


Discussion 


In a cladistic study of the tribe Kingsle- 
yini, Rodriguez & Pereira (1992) concluded 
that in Eudaniela pestai the exognath of the 
third maxilliped is less reduced than in oth- 
er members of the tribe. In addition, the ce- 
phalic lobe of the first male gonopod is 
formed by two separate plates which resem- 
ble an ancestral Strengeriana-type of gon- 
opod with three independent plates. For 
these reasons they considered “‘Eudaniela 
pestai aS approaching the ancestral pseu- 
dothelphusid condition”’’, a conclusion con- 
firmed by Rodriguez & Campos (1998). 
More recently, a cladistic analysis of mor- 
phological characters was conducted by 
Sternberg et al. (1999) to resolve the rela- 
tionships of the Eudaniela species complex. 
The result presented Eudaniela as a para- 
phyletic taxon, and E. pestai was positioned 
basal to the Eudaniela species complex, and 
to other representatives of the tribe King- 
sleyini, i.e. the genera Fredius Pretzmann, 
1967, Guinotia Pretzmann, 1965, Kingsleya 
Ortmann, 1897, and Microthelphusa Pretz- 
mann, 1968. In this study E. pestai is trans- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ferred to the new genus Achagua, and with 
the new species A. casanarensis the they 
seem to constitute a basal group from which 
Eudaniela species originated. Thus, Achag- 
ua likely represents a sister-group to all oth- 
er Kingsleyini. 


Acknowledgments 


I am especially grateful to G. Rodriguez 
for many ideas that substantially improved 
the manuscript. I am also indebted to R. 
Lemaitre for critically reading the manu- 
script. I thank J. Murillo for his helpful 
comments about cladistics. The illustrations 
were prepared by Juan C. Pinzon. 


Literature Cited 


Bott, R. 1970. Bemerkungen zu einigen Siisswasser- 
krabben (Crustacea: Decapoda).—Senckenber- 
giana Biologica 51(5/6):355-—361. 

Ortmann, A. 1897. Carcinologische Studien.—Zoolo- 
gische Jahrbticher, Abteilung fiir Systematik, 
Geographie and Biologie der Tiere 10:258—372. 

Pretzmann, G. 1965. Vorlaufiger Bericht tiber die Fam- 
ilie Pseudothelphusidae—Anzeiger der Oster- 
reichischen Akademie der Wissenschaften 
Mathematische Naturwissenschaftliche Klasse 
(1), 1:1-10. 

1967. Uber einige siidamerikanische Siis- 

swasserkrabben (Pseudothelphusidae). Vorlau- 

fige Mitteilung—Entomologische Nachrichten- 

blatt, Wien 14:23—26. 

1968. Neue siidamerikanische Stisswasser- 

krabben der Gattung Pseudothelphusa.—Ento- 

mologische Nachrichtenblatt, Wien 15:1—15. 

. 1971. Fortschritte in der Klassifizierung der 
Pseudothelphusidae.—Anzeiger der Mathema- 
tisch Naturwissenschaftliche der Osterreichisch- 
en Akademie der Wissenschaften (1)179(1—4): 
14-24. 

Rathbun, M. 1893. Descriptions of new species of 
American freshwater crabs.—Proceedings of 
the United States National Museum 16:649— 
661, pls. 73-77. 

Rodriguez, G. 1982. Les crabes d’eau douce 
d’Amérique. Famille des Pseudothelphusi- 
dae.—Faune Tropicale 22:1—223. 

, & M. R. Campos. 1998. A cladistic revision 

of the genus Fredius (Crustacea: Decapoda: 

Pseudothelphusidae) and its significance to the 

biogeography of the Guianan lowlands of South 

America.—Journal of Natural History 32:763— 

775. 

, & G. Pereira. 1992. New species, cladistic 


VOLUME 114, NUMBER 4 


relationships, and biogeography of the genus 
Fredius (Decapoda: Brachyura: Pseudothelphu- 
sidae) from South America.—Journal of Crus- 
tacean Biology 12:298-311. 

Smalley, A. 1964. A terminology for the gonopods of 
the American river crabs.—Systematic Zoology 
13:28-31. 


943 


Smithe, E B. 1975. Naturalist’s color guide-—The 
American Museum of Natural History, New 
York. Part 1: unnumbered pages. 

Sternberg von R., L. A. Galindo & E. M. Gonzalez. 
1999. Cladistic analysis of the Eudaniela spe- 
cies complex (Crustacea: Decapoda: Pseudoth- 
elphusidae).—Hydrobiologia 416:139—147. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(4):944—950. 2001. 


A new pontoniine shrimp of the genus Coralliocaris Stimpson, 1860 
(Crustacea: Decapoda: Palaemonidae) from the Ryukyu Islands 


Masako Mitsuhashi, Takahiro Fujino, and Masatsune Takeda 


(MM, MT) Department of Biological Sciences, Graduate School of Science, The University of 
Tokyo; Department of Zoology, National Science Museum, 3-23-1 Hyakunincho, Shinjuku-ku, 
Tokyo, 169-0073 Japan; 

(TF) Department of Biology, Faculty of Science, Yamagata University, 1-4-12 
Koshirakawa-machi, Yamagata, 990-8560 Japan 


Abstract.—A new species of coral-associated shrimp Coralliocaris tridens, 
is described from the Ryukyu Islands, southwestern Japan. The new species 
differs primarily from the eight congeneric species in having three teeth on the 
cutting edge of the dactylus of the second pereiopod. 


In 1969, the second author carried out a 
field trip to Ishigaki Island, the southern 
Ryukyu Islands, to investigate the coral reef 
shrimps of the family Palaemonidae. Most 
of the specimens collected were then iden- 
tified, but some remained unstudied. Re- 
cently, we re-examined the collection, and 
discovered two unusual specimens belong- 
ing to the pontoniine genus Coralliocaris 
Stimpson, 1860. The genus is composed of 
eight known species which are obligatory 
associates with scleractinian corals (Chace 
& Bruce 1993, Bruce 1998, Mitsuhashi 
2000). Detailed examination has shown that 
the two specimens represent an undescribed 
species described herein. 

The type specimens of the new species 
are deposited at the Kitakyushu Museum 
and Institute of Natural History, Kitakyushu 
(KMNH). The carapace length (CL), from 
the posterior margin of the orbit to the mid- 
point of the posterodorsal margin of cara- 
pace, is an indication of specimen size. The 
measurements and drawings were made 
with the aid of a drawing tube mounted on 
a LEICA MZ8 stereo microscope or Nikon 
70021 microscope. 


Coralliocaris tridens, new species 
Figs. 1-3 


Material examined.—Holotype: oviger- 
ous 2 (CL 3.05 mm), KMNH IvR 300002, 


Kabira Bay (24°26'N, 124°8’E), Ishigaki Is- 
land, Ryukyu Islands, southwestern Japan, 
from tabular coral (Acropora sp.), 28 Jul 
1969, coll. T. Fujino. Paratype: 1 sex undet. 
(CL 2.06 mm), KMNH IvR 300001, same 
data as holotype. 

Description of holotype.—Medium-sized 
shrimp, with typical shape for genus Cor- 
alliocaris. Body (Fig. 1) moderately de- 
pressed dorsoventrally. Rostrum (Figs. 1, 
2A) unarmed, rather abruptly narrowed in 
distal half, just reaching base of interme- 
diate segment of antennular peduncle; mid- 
rib well developed, broadened posteriorly; 
supraorbital eave nearly straight in dorsal 
view. Carapace (Figs. 1, 2A) glabrous; an- 
tennal spine submarginal; anterolateral an- 
gle broadly rounded. 

Abdomen (Figs. 1, 2A) with pleura of 
first 3 segments broadly expanded, forming 
marsupium; third segment feebly produced 
posterodorsally, partially covering lateral 
surfaces of fourth and fifth segments; fourth 
and fifth segments each with posteroven- 
trally rounded pleuron; sixth segment lon- 
ger than fifth segment, with acute postero- 
ventral tooth; posterolateral process blunt. 

Telson (Fig. 3A) elongate, subtriangular, 
length about 3 times greatest width; dorsal 
surface with 2 pairs of small spines dorso- 
laterally, anterior pair situated slightly pos- 


VOLUME 114, NUMBER 4 


945 


Fig. 1. 
Habitus, dorsal. Scale: 3 mm. 


terior to midlength of telson, posterior pair 
situated at about midlength between ante- 
rior pair and posterior margin; posterior 
margin rounded, with 3 pairs of spines, lat- 
eral pair shortest; intermediate pair longest, 
stout; submedian pair somewhat shorter 
than intermediate; simple fine seta arising 
from near bases of submedian and inter- 
mediate pairs of spines. 

Eyes (Figs. 1, 2A) moderately large; eye- 
stalk subcylindrical, 1.4 times as long as 
wide; cornea oval, slightly inflated. 

Antennular peduncle (Fig. 1) reaching 
0.7 of scaphocerite; basal segment broad, 
greatest width approximately equal to 
length of medial margin; ventral surface 
with small tooth at posterior 0.7; outer mar- 
gin of stylocerite convex along general out- 
line of basal segment, reaching level of dis- 
tal 0.2 of medial margin. Intermediate seg- 
ment as long as wide, with row of long plu- 
mose setae on medial margin. Distal 
segment weakly broadened distally; upper 
flagellum biramous, with proximal fused 
part composed of 7 articles. 

Basicerite of antenna with prominent lat- 
eral spine; scaphocerite twice as long as 
broad, lateral margin straight, armed with 
stout tooth distally. 

Third maxilliped with stout endopod; ba- 


Coralliocaris tridens, new species. Holotype (KMNH IvR 300002) ovigerous female (CL 3.05 mm). 


sis and ischiomeral segments incompletely 
fused; ischiomeral segment 3 times as long 
as basis, unarmed on lateral margin; carpus 
swollen medially, 1.5 times broader than 
terminal segment, with numerous, fine pul- 
mose setae on medial surface; terminal seg- 
ment short, teardrop-shaped, with dense 
plumose setae on dorsal surface; exopod al- 
most reaching distal end of endopod; 
rounded epipod present; arthrobranch with 
6 lamellae. 

First pereiopod (Fig. 3B, C) slender, ex- 
ceeding distal margin of scaphocerite by 
length of chela and half of carpus. Chela 
about half as long as carpus, gradually ta- 
pering distally; fingers half length of palm, 
each with entire cutting edge; palm with 
about 10 transverse rows of anteriorly 
curved, serrate setae on ventral surface. 
Carpus gradually widened distally, ventro- 
distally with transverse row of setae. Merus 
3 times longer than ischium. 

Second pereiopods (Fig. 1) subequal in 
length and shape. Chela large, elongate, 
somewhat compressed laterally. Dactylus 
about half length of palm, nearly straight 
in ventral view, weakly curved and distally 
hooked in lateral view, lacking longitudi- 
nal ridge on lateral face; cutting edge with 
3 subtriangular equidistant teeth on proxi- 


946 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


F H - 


Fig. 2. Coralliocaris tridens, new species. A, holotype (KMNH IvR 300002) ovigerous female (CL 3.05 
mm); B—H, paratype (KMNH IvR 300001), sex undet. (CL 2.06 mm). A, body, lateral, cephalic and thoracic 
appendages omitted; B, antennular peduncles and anterior part of rostrum, dorsal; C, left mandible, ventral; D, 
left maxillula, ventral, lower lacinia missing; E, left maxilla, ventral; E left first maxilliped, ventral; G, left 
second maxilliped, ventral, part of epipod missing; H, left third maxilliped, ventral. Scales: 1 mm. 


mal half. Fixed finger with 4 teeth, distal dactylus on slightly medial to cutting edge. 
3 similar to dactylar teeth, proximal tooth Palm with weakly concave lateral profile 
faintly tridenticulate; 3 shallow concavities in dorsal view. Carpus short, about 0.3 as 
fitting to 3 teeth on opposable margin of long as palm, cup-shaped; distal margin 


VOLUME 114, NUMBER 4 947 


Fig. 3. Coralliocaris tridens, new species. A—C, H, I, holotype (KMNH IvR 300002) ovigerous female (CL 
3.05 mm); D-G, J, paratype (KMNH IvR 300001), sex undet. (CL 2.06 mm). A, telson, dorsal; B, left first 
pereiopod, ventral; C, chela of right first pereiopod, lateral; D, left second pereiopod, ventral; E, fingers, same, 
medial; EK left third pereiopod, lateral; G, same, dactyl and distal part of propodus, medial; H, exopod of right 
uropod, dorsal; I, J, lateral part of exopod of right uropod, dorsal. Scales: 1 mm, except C and G, 0.5 mm. 


with a stout process on ventral part, ob- tooth. Ischium compressed, about half 
scurely denticulate dorsally. Merus about length of merus. 

half length of palm and twice as long as Third to fifth pereiopods similar, robust. 
carpus, with moderately strong distomedial Dactyli short, distally blunt, each with 


948 


strong acute unguis on midlength of margin 
of extensor surface. Propodi 4 times longer 
than wide, with transverse rows of dense 
curly setae distally. Carpi short, about half 
of propodi; meri subequal in length to pro- 
podi and about twice as long as carpi, 
slightly narrowed distally. Ischia about half 
as long as meri. 

Pleopods normal; first pleopod without 
setae on surface of protopod; endopod sub- 
cylindrical, elongate, overreaching half of 
exopod. 

Uropod (Fig. 3H, I) overreaching tip of 
telson; exopod with acute immovable tooth 
and 2 movable spines at distal 0.3 of lateral 
margin on both sides. 

About 70 ova at early-eyed stage present, 
measuring 0.64—0.74 mm X 0.48—0.53 mm. 

Description of paratype.—Body more 
slender than holotype. 

Rostrum (Fig. 2B) extending to mid- 
length of distal segment of antennular pe- 
duncle. Abdominal segments more slender 
than holotype, with feebly developed pleu- 
ra. Antennular peduncle (Fig. 2B) with 
well-developed stylocerite; tip of stylocerite 
protruding obliquely forward from _ basal 
segment leaving v-shaped notch. 

Mandible (Fig. 2C) deeply divided in in- 
cisor and molar processes, without palp; 
molar process slender with dense bristles 
distally; incisor process tapering distally, 
armed with 4 teeth on distomedial margin, 
intermediate 2 teeth somewhat smaller than 
outer teeth. Maxillula (Fig. 2D) with short 
apical seta on tip of inner lobe of palp; up- 
per lacinia with several stout setae on distal 
margin. Maxilla (Fig. 2E) with short, sim- 
ple endite bearing long apical seta; palp 
slender, feebly tapering, twice as long as 
endite. First maxilliped (Fig. 2F) with short 
non-setose palp; basial endite rounded, 
fringed with setae on medial margin, with- 
out distinct notch separating it from coxal 
endite; coxal endite with a few setae at me- 
dial margin; exopod approximately 3 times 
as long as basial endite, with several plu- 
mose setae distally; caridean lobe short, 
broad. Second maxilliped (Fig. 2G) typical 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


for genus; dactylar segment with dorsally 
curved setae on medial margin; propodus 
not produced anteriorly, with several setae 
on medial margin. Third maxilliped (Fig. 
2H) similar to that of holotype. 

First pereiopod stouter than in holotype. 
Chela 0.6 times as long as carpus. Second 
pereiopod (Fig. 3D, E) similar to that of 
holotype on left side, but smaller on right 
side, right chela about half as long as left 
chela. Cutting edge of fixed finger of left 
chela (Fig. 3E) with most proximal tooth 
truncate, with 2 faintly developed denticles; 
dactylus of small right chela with small 
tooth and fixed finger with 2 small teeth. 
Third pereiopod (Fig. 3E G) to fifth pereio- 
pod slightly more slender than in holotype; 
propodi with dense setae distally, as in ho- 
lotype. 

Pleopods without specific features; en- 
dopod and exopod of first pleopod short, 
narrow triangular. 

Posterior half of left uropod broken; right 
exopod (Fig. 3J) with immovable tooth and 
1 spine on lateral margin. 

Coloration.—The preserved specimens 
in spirit are uniformly whitish-cream. Color 
in life not recorded. 

Etymology.—The specific name is a 
combination of the Latin, tri (=three) and 
dens (=tooth), in reference to the charac- 
teristic three teeth on the cutting edge of the 
dactylus of the second pereiopod. 

Distribution.—Known so far only from 
Kabira, Ishigaki Island, Ryukyu Islands. 

Host.—tThe host coral is an unidentified 
species of Acropora, like in most of Cor- 
alliocaris species (Bruce 1972, 1977). 

Remarks.—The sex of the paratype could 
not be determined, because the appendix 
masculina is not differentiated. 

Coralliocaris tridens, new species, is 
readily distinguished from the other species 
of Coralliocaris by having three teeth on 
the cutting edge of dactylus of the second 
pereiopod. The cutting edge of the dactylus 
of the second pereiopod is armed with two 
teeth in C. brevirostris Borradaile, 1898, C. 
nudirostris (Heller, 1861) and C. venusta 


VOLUME 114, NUMBER 4 


Kemp, 1922, and with one blunt projection 
in C. superba Dana, 1852 and C. taiwanen- 
sis Fujino & Miyake, 1972. In C. graminea 
Dana, 1852, C. viridis Bruce, 1974 and C. 
macrophthalma (H. Milne Edwards, 1837), 
the cutting edge of the dactylus of the sec- 
ond pereiopod is unarmed, bearing a fossa 
in which fits a flattened tooth on the op- 
posable margin (Chace & Bruce 1993, 
Bruce 1998, Mitsuhashi 2000). The arma- 
ture of the dactylus is constant and without 
doubt one of the important character to dis- 
tinguish the species. 

Coralliocaris tridens is most similar to 
C. nudirostris in having unarmed rostrum, 
triangular teeth on the cutting edges of fin- 
gers of the second pereiopods, and straight 
subdistal part at the cutting edges of the 
fixed finger of the second pereiopod. We 
fortunately could examine the type speci- 
mens of C. nudirostris in the collections of 
the Natural History Museum in Vienna and 
found that the dactylus of the second pe- 
reiopod bears a longitudinal ridge on its 
medial surface, and differs from that of C. 
tridens, in which the medial surface is not 
ridged and smooth (Fig. 3E). 


Acknowledgments 


We are very grateful to Y. Yabumoto, Ki- 
takyushu Museum and Institute of Natural 
History, for making available specimens. 
Thanks are also expressed to M. Tiirkay, 
Senckenberg Natural History Museum, and 
P. C. Dworschak, Natural History Museum, 
Vienna, for information on Heller’s collec- 
tion and loan of type specimens, respec- 
tively. We are deeply indebted to T. Komai, 
Natural History Museum and Institute, Chi- 
ba, and A. J. Bruce, Queensland Museum, 
Australia, for suggestions to the manuscript. 
We also thank R. Lemaitre, National Mu- 
seum of Natural History, Smithsonian In- 
stitution; A. Anker, of Muséum national 
d’ Histoire naturelle, Paris; and an anony- 
mous reviewer for valuable comments. This 
study was partly supported by the Sasagawa 


949 


Scientific Research Grant from the Japan 
Science Society. 


Literature Cited 


Borradaile, L. A. 1898. A revision of the Pontoni- 
idae.—Annals and Magazine of Natural Histo- 
ry, ser. 7, 2:376—391. 

Bruce, A. J. 1972. A review of information upon the 
coral hosts of commensal shrimps of the sub- 
family Pontoniinae, Kingsley, 1878 (Crustacea, 
Decapoda, Palaemonidae). Pp. 399—417 In Pro- 
ceedings of the Symposium on Corals and Coral 
Reefs. The Marine Biological Association of In- 
dia. 

. 1974. Coralliocaris viridis sp. nov., a prelim- 

inary note (Decapoda, Natantia, Pontoni- 

inae).—Crustaceana 26:222—224. 

. 1977. The hosts of the coral-associated Indo- 

West-Pacific pontoniine shrimps.—Atoll Re- 

search Bulletin 205:1—19. 

. 1998. New keys for the identification of Indo- 
West Pacific coral associated pontoniine 
shrimps, with observations on their ecology 
(Crustacea: Decapoda: Palaemonidae).—Ophe- 
lia 46:29—46. 

Chace, F A. Jr, & A. J. Bruce. 1993. The caridean 
shrimps (Crustacea: Decapoda) of the Albatross 
Philippine Expedition, 1907-1910, part 6: Su- 
perfamily Palaemonoidea.—Smithsonian Con- 
tributions to Zoology 543:1—152. 

Dana, J. D. 1852. Conspectus Crustaceorum quae in 
Orbis Terrarum circumnavigatione, Carolo 
Wilkes e Classe Republicae Foederatae Duce, 
lexit et descripsit—Proceedings of the Acade- 
my of Natural Sciences of Philadelphia 6:10— 
28. 

Fujino, T., & S. Miyake. 1972. A new pontoniinid 
shrimp of the genus Coralliocaris Stimpson 
from Taiwan (Crustacea, Decapoda, Pontoni- 
inae).—Occasional Papers of Zoological Labo- 
ratory Faculty of Agriculture Kyushu Univer- 
sity 3:91—98. 

Heller, C. 1861. Synopsis der im rothen Meere vor- 
kommenden Crustaceen.—Verhandlungen der 
Kaiserlich-k6niglichen Zoologisch-Botanischen 
Gesellschaft in Wien 11:1—32. 

Kemp, S. 1922. Notes on Crustacea Decapoda in the 
Indian Museum, XV: Pontoniinae.—Records of 
the Indian Museum 24:113-—288. 

Milne Edwards, H. 1837. Histoire naturelle des Crus- 
tacés, comprenant l’anatomie, la physiologie et 
la classification de ces animaux 2:1—532. Paris 

Mitsuhashi, M. 2000. A revision of the genus Coral- 
liocaris Stimpson, 1860 (Crustacea: Decapoda: 
Palaemonidae) from Japan.—I. O. P. Diving 
News 11():2-7. (In Japanese with English 
summary) 


950 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Stimpson, W. 1860. Crustacea Macrura. Pars VIII of 
Prodromus descriptionis animalium evertebra- 
torum, quae in Expeditione ad Oceanum Paci- 
ficum Septentrionalem, a Republica Federata 


missa, Cadwaladaro Ringgold et Johanne Rodg- 
ers Ducibus, observavit et descripsit.—Proceed- 
ings of the Academy of Natural Sciences of 
Philadelphia 12:22—47. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
114(4):951—963. 2001. 


A new genus and a new species of crab of the family Goneplacidae 
MacCleay, 1838 (Crustacea: Decapoda: Brachyura) from the tropical 
western Atlantic 


Rafael Lemaitre, Julio Garcia-Gomez, Richard v. Sternberg, and Néstor H. Campos 


(RL) Department of Systematic Biology, National Museum of Natural History, Smithsonian 
Institution, Washington, D.C. 20560-0163, U.S.A., e-mail: lemaitre.rafael@nmnh.si.edu; 
(JGG) Division of Marine Biology and Fisheries, Rosenstiel School of Marine and Atmospheric 
Science, University of Miami, 4600 Rickenbacker Causeway, Miami, Florida 33149, U.S.A.; 
(RvS) Department of Systematic Biology, National Museum of Natural History, Smithsonian 
Institution, Washington, DC 20560-0163, U.S.A.; current address: Taxonomy Project, National 
Center for Biotechnology Information (GenBank), National Institutes of Health, Building 45, 
Room 6An.18D-30, Bethesda, Maryland, U.S.A.; 

(NHC) Universidad Nacional de Colombia, c/o Instituto de Investigaciones Marinas y Costeras, 
Apartado Aéreo 1016, Santa Marta, Colombia 


Abstract.—A new monotypic genus, Nancyplax, and a new species, N. vossi, 
of the family Goneplacidae MacLeay, are described from the southern Carib- 
bean and northeastern coast of South America. Characters derived from male 
pleopods 1 and 2, and male abdomen-sternoabdominal cavity, allow Nancyplax 
to be assigned to the Euryplacinae Stimpson. The carapace in N. vossi, how- 
ever, strongly resembles in overall conformation some species in the Carci- 
noplacinae H. Milne Edwards. In addition, NV. vossi has a well developed strid- 
ulatory organ, a feature shared with only one other euryplacine species, some 
carcinoplacines, and some species in the Goneplacinae MacLeay. The mosaic 
habitus of Nancyplax precludes resolution of relationships with other eurypla- 
cine genera. Morphological similarities of N. vossi with species of Carcinoplax 
H. Milne Edwards, Psopheticus Wood-Mason, Psopheticoides Sakai, and Tri- 
zocarcinus Rathbun, are summarized. 


The family Goneplacidae MacLeay, 1838 
currently contains numerous problematic 
genera with xanthoid affinities that defy 
phylogenetic arrangement. Guinot (1969a, 
1969b, 1969c) proposed a subfamilial ar- 
rangement which subsequently has been 
modified by the removal and addition of 
subfamilies (Manning & Holthuis 1981, 
Guinot 1986, Ng & Wang 1994, Hendrickx 
1998). As a result, the Goneplacidae is now 
generally divided by carcinologists into six 
subfamilies: Carcinoplacinae H. Milne Ed- 
wards, 1852, Chasmocarcininae Seréne, 
1964, Euryplacinae Stimpson, 1871, 
Goneplacinae MacLeay, 1838, Pseudozi- 
inae Alcock, 1898, and Trogloplacinae 


Guinot, 1986. Of these six subfamilies, the 
Carcinoplacinae, Euryplacinae, and Gone- 
placinae are considered closely related 
(Guinot 1969a, 1969b, 1969c). The remain- 
ing three subfamilies appear to have no 
close affinities either among themselves or 
with other subfamilies of Goneplacidae, but 
should remain in the family until detailed 
phylogenetic analyses are conducted. 
Although in-depth taxonomic reviews 
have been completed for goneplacid genera 
such as Carcinoplax H. Milne Edwards, 
1852 (see Guinot 1989), and Psopheticus 
Wood-Mason, 1892 (see Guinot 1990), the 
systematic position and relationships of 
many taxa remain to be fully evaluated. Ob- 


952 


stacles to resolving relationships include 
vague subfamilial definitions and mosaic 
taxonomic distributions of character states. 
Furthermore, new forms which often exhib- 
it puzzling combinations of generic-level 
features continue to be discovered through- 
out the world oceans. 

In the last 25 years a substantial number 
of new taxa from the Atlantic and Pacific 
coasts of the American continents have 
been added to the Goneplacidae (e.g., Go- 
mez & Ortiz 1975, Hernandez Aguilera 
1982, Guinot 1984, Garth 1986, Hendrickx 
1989, 1998; Vazquez-Bader & Gracia 
1991, 1995; Tavares 1996). Recent exami- 
nation of numerous unstudied specimens 
deposited in the collections of the Rosen- 
stiel School of Marine and Atmospheric 
Science, University of Miami, and obtained 
in the 1960s during deep-sea cruises of the 
R/V Pillsbury to the Atlantic coast of north- 
ern South America, revealed the existence 
of yet another undocumented goneplacid 
crab of unclear affinities that requires as- 
signment to a new genus and species. Spec- 
imens of this same crab were also recently 
collected in 1995 from the Caribbean coast 
of Colombia. Herein, a new monotypic ge- 
nus and new species are described for this 
goneplacid crab. 


Material and Methods 


Specimens used were collected in 1968 
and 1969 during investigations of the ma- 
rine fauna of the tropical Atlantic on board 
the R/V Pillsbury by the then Institute of 
Marine Sciences, University of Miami, now 
Rosenstiel School of Marine and Atmo- 
spheric Science (RSMAS); and in 1995 
during a joint expedition along the Carib- 
bean coast of Colombia on board the re- 
search vessel B/I Ancon, conducted by staff 
from the Colombian Navy’s Centro de In- 
vestigaciones Oceanograficas e Hidrografi- 
cas, Cartagena (CIOH), Instituto de Inves- 
tigaciones Marinas y Costeras, Santa Marta 
(INVEMAR), and National Museum of 
Natural History, Smithsonian Institution, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Washington, DC (USNM). Sampling meth- 
ods used on the R/V Pillsbury can be found 
in Voss (1966), and on the B/I Ancén in 
Lemaitre & Campos (2000). The holotype 
has been deposited in the USNM, with par- 
atypes in the reference collection of IN- 
VEMAR, and the Invertebrate Museum of 
RSMAS (UMML). In general, the descrip- 
tive terminology used follows Guinot 
(1969a, 1969b, 1969c, 1989, 1990). The 
term “‘press-button”’ is used as defined by 
Guinot & Bouchard (1998) for the abdom- 
inal locking mechanism. For the stridulat- 
ing organ, the terms “pars stridens” and 
““plectrum”’ are used in accordance with 
Guinot-Dumortier & Dumortier (1960) and 
Guinot (1990). In the “‘Material examined”’ 
section, measurements listed are in milli- 
meters (mm), given in the order: total car- 
apace length (CL) X carapace width (CW); 
and abbreviations are as follows: ovig, 
ovigerous; P, R/V Pillsbury; sta, station. 


Nancyplax, new genus 


Diagnosis.—Carapace roughly hexago- 
nal in adults, distinctly broader than long; 
dorsal surface convex, with depression on 
each side of juncture between gastric and 
cardiac regions. Branchial region with 
rounded, low mesial protuberance and 
prominent, granular metabranchial tubercle. 
Exorbital angle indistinct, consisting of a 
low, rounded lobe; anterolateral teeth ab- 
sent except for epibranchial tooth. Epibran- 
chial (lateral) spine pronounced, sharp. 
Front straight, gently deflexed, double-mar- 
gined. Orbits oblique; medial angle of in- 
fraorbital margin with well-developed oc- 
clusive lobe. Antennules folding transverse- 
ly. Suture separating subhepatic and 
pterygostomial regions flanked on ptery- 
gostomial region by longitudinal ridge with 
row of small tubercles. Stridulating mech- 
anism present, well developed. 

Lateral margins of buccal frame strongly 
divergent anteriorly. No endostomial ridges 
present. Posterior margin of epistome with 
deep medial notch and 2 incisions on either 


VOLUME 114, NUMBER 4 


side extending to anterior margin. Interca- 
lary apophysis between basal antennal ar- 
ticle and pterygostomial border well devel- 
oped and subrectangular in outline. Third 
maxillipeds not completely covering buccal 
frame; merus with mesial margin convex, 
distolateral angle forming rounded lobe, 
distal margin concave. 

Thoracic sternum width between lateral 
margins of fifth episternites about 1.30 
times greater than total length. Sternite 1 
separated from sternite 2 by indistinct sul- 
cus. Anterior border of sternite 3 much wid- 
er than posterior margin of sternite 2. Ster- 
nites 3 and 4 demarcated by shallow rem- 
nants of sulcus confined to lateral regions. 
Sulci separating sternites 4 and 5, 5 and 6, 
and 6 and 7, interrupted medially. Sulcus 
delimiting sternites 7 and 8 complete. Me- 
dian septum of endosternite 8 externally 
visible as median, longitudinal sulcus. 
Press-button knobs on fifth sternite distinct, 
adjacent to sternite 4. Episternites 4 through 
7 developed and broadly lobate posteriorly. 
Sternoabdominal cavity deep, triangular in 
outline. Episternite 7 covering small portion 
of male seminal duct aperture on coxa of 
pereopod 5 and part of short seminal groove 
on sternite 8. 

Abdomen in both sexes with second seg- 
ment leaving small portion of episternite 8 
visible. Male abdomen triangular in overall 
outline, with 6 free articulating segments 
and telson; third segment overlapping part 
of coxae of fifth pereopods and completely 
covering gonopores located at base of cox- 
ae; telson triangular, with distal margin 
rounded. 

Male with pleopod 1 sinuous, much 
broader at base than at apex, tapering to 
very slender tip, spinose; euryplacine in or- 
ganization (cf. Guinot 1969b). Pleopod 2 
short; apical lobe reduced and separated 
from subterminal segment by incomplete 
fissure. 

Type species.—Nancyplax vossi, new 
species, by present designation. 

Gender.—Feminine. 

Etymology.—The name of the genus is in 


953 


honor of Mrs. Nancy Voss, Research Pro- 
fessor at the Rosenstiel School of Marine 
and Atmospheric Science, University of 
Miami. The genus name is formed by com- 
bining her first name with the Greek suffix 
plax, meaning flat (although morphologi- 
cally inappropriate here for the only species 
in the genus) and commonly used in nam- 
ing genera of the Goneplacidae. Mrs. N. 
Voss has for many years vigorously pro- 
moted the study of the invaluable tropical 
deep-sea collections housed in her institu- 
tion. 

Remarks.—Nancyplax, new genus, can 
be placed in the Euryplacinae with relative 
confidence. This placement is based first on 
the structure of male pleopods 1 and 2. Un- 
like the pleopods 1 seen in the Carcinopla- 
cinae and Goneplacinae, euryplacine pleo- 
pods 1 are somewhat broad at the base, al- 
most uniformly tapering to a very slender 
apex with patches of distinct spines on the 
surface (Guinot 1969b). This particular ple- 
opod | morphology is probably a defining 
characteristic of the Euryplacinae. In con- 
trast, pleopods 1 of carcinoplacines and go- 
neplacines are considerably stouter along 
the distal half of the terminal segment, and 
the apex may or may not possess one or 
more distal lobes (Guinot 1969b, 1989, 
1990). Second, pleopod 2 is short in Nan- 
cyplax with a rudimentary apical lobe as it 
is in all other euryplacine taxa (Guinot 
1969b); carcinoplacines and goneplacines 
both have long second pleopods with well 
defined flagella, and a specific apical con- 
formation (Guinot 1969b, 1989, 1990). 
Third, Nancyplax exhibits a male abdomen 
outline and sternoabdominal cavity organi- 
zation that is very close to those seen in 
euryplacines such as species of Trizocar- 
cinus Rathbun, 1914, e.g., T. dentatus 
(Rathbun, 1893). These characters permit 
Nancyplax to be excluded from either the 
Carcinoplacinae or Goneplacinae. 

The carapace in the single species of 
Nancyplax is nevertheless unlike that of any 
other euryplacine. In terms of overall shape 
the carapace strongly resembles those seen 


954 


in some species of Psopheticus and Carci- 
noplax, especially C. longispinosa Chen, 
1984. However, Psopheticus species have 
in common with Nancyplax vossi the pres- 
ence of a stridulatory organ (Guinot-Du- 
mortier & Dumortier 1960, Guinot 1990), 
whereas this organ is absent from members 
of Carcinoplax. Within the Euryplacinae, 
Trizocarcinus dentatus also has a well de- 
veloped stridulatory organ, and the ptery- 
gostomial ridges are identical in form to 
those of the single species of Nancyplax, 
although the two species have different car- 
apace outlines. Species of the goneplacine 
Ommatocarcinus White, 1852 and the car- 
cinoplacine Bathyplax A, Milne-Edwards, 
1880, likewise possess a Nancyplax-like 
stridulatory organ, but here again carapace 
morphology is not equivalent among the 
three genera. Given the patchy taxonomic 
distribution of the stridulatory organ in go- 
neplacids, and the disparate carapace shapes 
observed in some genera such as Carcino- 
plax (see Guinot 1989), it would appear that 
characters from the carapace cannot be used 
to argue against placing Nancyplax in the 
Euryplacinae. 

The morphology of Nancyplax exempli- 
fies many of the problems inherent in go- 
neplacid systematics. To partially reiterate, 
this new genus can be said to have a mosaic 
habitus. The male pleopods, male abdomen 
outline, and sternoabdominal cavity confor- 
mation are euryplacine. The shapes of the 
male telson and third maxilliped merus are 
very close to those in Trizocarcinus den- 
tatus. Nancyplax has the same type of strid- 
ulatory organ as seen in 7. dentatus and 
some carcinoplacines. Nancyplax and Car- 
cinoplax longispinosa have almost equiva- 
lent carapace anatomies except for the strid- 
ulatory ridges. The lack of defined grooves 
on episternite 8 for the seminal ducts sug- 
gests that Nancyplax is not as derived as 
many other carcinoplacines, euryplacines, 
and goneplacines (Guinot 1969b). For these 
reasons, it is unclear whether Nancyplax is 
a basal euryplacine or alternatively, a high- 
ly modified taxon. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Nancyplax vossi, new species 
Figs. 1—5 


Material examined.—Holotype: 3 10.7 
x 15.7 mm, off Venezuela, sta P-752, 
11°06.3'N, 68°14.6'W, 95-132 m, 26 July 
1968, USNM 308995. 

Paratypes: 1 ¢ 13.9 X 20.2 mm, USNM 
308996; 2 ovig 2 8.6 X 12.7 mm, 10.6 X 
15.2 mm, UMML 32.8776; off Venezuela, 
sta P-752, 11°06.3'N, 68°14.6'W, 95-132 
m, 26 Jul 1968.—2 6 4.5 X 5.8 mm, 14.3 
x 20.1 mm, USNM 308993; 1 3 13.0 xX 
19.4 mm, INVEMAR-CRU 2926; SW Isla 
Rosario, Islas del Rosario, Colombia, “‘Ex- 
pedici6n CIOH-INVEMAR-Smithsonian’’, 
sta T18, 10°11.97"N, 75°53.41’W, 150-155 
m, 7 Aug 1995.—2 ¢ 7.9 X 11.7 mm, 9.2 
x 13.4 mm, off SE coast of Trinidad, sta 
P-837, 10°09.8'N, 60°34.3'W, 55 m, 30 Jun 
1969, UMML 32.9079.—4 ¢ 6.0 X 7.6 
mm, 7.5 X 10.4 mm, 7.8 X 10.7 mm, 8.5 
X 12.1 mm, 1 2 6.0 X 8.0 mm, off SE 
coast of Trinidad, sta P-836, 9°56.5’N, 
60°46"W, 57-59 m, 30 Jun 1969, UMML 
32.9078.—1 3 12.1 X 17.2 mm, 1 2 11.9 
x 17.5 mm, USNM 308994; 1 ¢ 11.0 X 
15.7 mm, 2 2 10.7 X 16.0 mm, 11.3 X 16.0 
mm, UMML 32.8775; off Surinam, sta P- 
671, 7°07'N, 55°08’W, 64 m, 11 Jul 1968. 

Diagnosis.—Carapace about 0.65 to 0.75 
as long as broad, smooth, glabrous; regions 
weakly demarcated. Front straight, at most 
about 0.40 as long as CW; frontal lobes di- 
vided by weak median cleft. Anterolateral 
margins with strong epibranchial (lateral) 
spine. Pterygostomian region with well de- 
veloped stridulating ridge. Male chelae gla- 
brous; female chelae with dense fringe of 
setae on dorsal and ventral margins. Dactyls 
of pereopods 2—5 each with 3 longitudinal 
fringes of setae dorsally and 1 fringe ven- 
trally. Male pleopod 1 very slender distally, 
with medial shoulder densely spinose, and 
minute well-spaced spines on distal third. 

Description.—Carapace (Figs. 1, 3a) 
subhexagonal, about 0.65 to 0.75 as long as 
broad; dorsal surface convex, superficially 
smooth and glabrous, microscopically with 


VOLUME 114, NUMBER 4 


955 


Fig. 1. 


numerous punctae medially on gastric, car- 
diac and intestinal regions, and numerous 
minute granules on hepatic and branchial 
regions. Regions indistinctly demarcated; 
gastric and hepatic regions evenly convex; 
with distinct depression on each side of 
juncture between gastric and cardiac re- 
gions; branchial regions each with low, 
rounded protuberance medially, and dis- 
tinct, granular metabranchial tubercle. Front 
(Figs. 2a, 3b) straight, about 0.30 (adults) 
to 0.40 (young) times as long as CW; sep- 
arated on each side from preorbital lobe by 
small notch; with double margin consisting 
of lower and upper, pseudofrontal margins, 
lower margin extending slightly beyond 
pseudofrontal; pseudofrontal margins divid- 
ed into lobes by weak medial cleft (weaker 


Nancyplax vossi, new species, paratype male 14.3 X 20.1 mm, SW of Isla Rosario, Islas del Rosario, 
Colombia, sta T18, USNM 308993: dorsal view. 


on lower margin); lateral, inferior lobe pro- 
duced and extending ventrally to third an- 
tennal segment near articulation between 
the second and third segments. Anterolat- 
eral margins broadly rounded, lacking any 
indication of teeth or spines except for 
prominent, strong epibranchial (lateral) cor- 
neous-tipped spine directed slightly for- 
ward in dorsal view and slightly upward in 
frontal view (Figs. 1, 2a). Epibranchial (lat- 
eral) spine in adults nearly as long as orbital 
length. Posterolateral and posterior margins 
broadly rounded. Subhepatic region mi- 
nutely granulose. Pterygostomian region 
(Fig. 2a) smooth, with well developed, 
raised stridulating ridge (pars stridens) with 
numerous transverse striae bordering suture 
between pterygostomian and subhepatic re- 


956 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. 


Nancyplax vossi, new species, a, c, male 13.9 X 20.2 mm, sta P-752, off Venezuela, USNM 308996; 


b, male 14.3 X 20.1 mm, SW of Isla Rosario, Islas del Rosario, Colombia, sta T18, USNM 308993. a, carapace 
and cephalic appendages, including basal segments and part of merus of left cheliped, frontal view; b, abdominal 
segments 1 to 3, coxae 4 and 5, and episternite 7, and visible part of episternite 8, ventral view; c, sternites 7 
and 8, episternites 6 and 7, coxae 4 and 5, left side (abdomen removed), ventral view. Abbreviations: al-3, 
abdominal segments 1—3; c, posterior margin of carapace; cx4,5, coxae of pereopods 4 and 5; ep6-8, episternites 
6 to 8; g, gonopore; ol, occlusive lobe; pl, plectrum; ps, par stridens; s7,8, sternites 7 and 8. Solid black indicates 


membranes. Scales equals 2 mm (a, b), and 1 mm (c). 


gions. Ventral, pterygostomial border of 
carapace flanking coxae of pereopods 1 
concave, lined with setae on inner margin. 

Orbits (Figs. 2a, 3a, b), about 0.50 
(adults) to 0.60 (young) as long as front, 
margins minutely granulose, with small, 
blunt pre and postorbital lobes; supraorbital 
margin with 2 short sutures and fringe of 
well-spaced setae; infraorbital margin ex- 
posed in dorsal view, with short suture just 
below postorbital lobe and subrectangular 
occlusive lobe. Eyestalks short, completely 
fitting in orbits when retracted; corneae 
slightly dilated; peduncles sparsely setose. 

Antennules folding transversely into 
deep fossae, segments similar in length; 


basal segment considerably inflated and oc- 
cupying nearly entire antennular fossae, 
penultimate and ultimate segments slender. 
Antennae with first segment immovable, 
short, semicircular and enclosing urinary 
opening; second to fourth segments movy- 
able, longer than broad; flagellum over- 
reaching antennules when fully extended. 
Third maxillipeds (Fig. 3c) with bases 
separated by triangular tip of thoracic ster- 
num. Ischium longer than broad; outer sur- 
face with median longitudinal furrow; me- 
sial margin with row of teeth and fringe of 
setae. Merus about as long as broad; dis- 
tolateral angle produced into lobe with mi- 
nutely granular outer surface and marginal 


VOLUME 114, NUMBER 4 


Fig. 3. 


957 


PDD pip 


Nancyplax vossi, new species. a, young male 4.5 X 5.8 mm, SW of Isla Rosario, Islas del Rosario, 


Colombia, sta T18, USNM 308993: b, d, f, male 13.9 X 20.2 mm, sta P-752, off Venezuela, USNM 308996; 
c, male 14.3 X 20.1 mm, sta T18, USNM 308993; e, female 9.5 X 13.4 mm, sta T18, USNM 308993. a, 
carapace, eyes and antennae, dorsal view; b, frontal region, eyes and antennae, dorsal view; c, third right 
maxilliped, outer view; d, left and right chelae male, outer view; e, left and right chelae of female, outer view; 
f, merus and carpus of right cheliped, dorsal view. Abbreviation: pl, plectrum. Scales equal 1 mm (a-c, e, f), 


and 2 mm (d). 


setae; outer mesial surface minutely gran- 
ular, setose. Palp about as long as ischium, 
marginally setose. Exopod with basal seg- 
ment reaching to tip of distolateral angle of 
merus; mesial surface minutely granulose. 
Chelipeds sexually dimorphic. Male che- 
lae (Fig. 3d) slightly unequal in size right 


from left, mostly glabrous. Dactyl about as 
long as palm. Fingers not leaving gap when 
closed, each terminating in inwardly curved 
corneous claw; outer surface minutely 
punctate; cutting edges consisting of row of 
small sharp or blunt calcareous teeth inter- 
spersed with larger calcareous teeth. Palm 


958 


glabrous; dorsal margin broadly curved, 
ventral margin nearly straight. Carpus (Fig. 
3f) with outer surface sparsely granulose, 
setose; inner margin armed proximally with 
strong spine curving inward distally and 
terminating in corneous tip. Merus (Fig. 3f) 
granulose except on dorsal face and plec- 
trum; with sparse row of setae on dorsome- 
sial margin; inner, proximal margin with 
stridulating ridge (plectrum). 

Female chelipeds subequal, differing 
from those of male as follows: chelae (Fig. 
3e) densely setose on dorsal and ventral 
margins; dactyl about 1.40 times as long as 
palm; fixed fingers and palm with dorsal 
and ventral surfaces granulose; carpus with 
outer face densely setose. 

Pereopods 2—5 (Fig. 4) slender; segments 
with lateral and mesial faces glabrous. Pe- 
reopods 2—4 subequal in length, about 1.60 
times as long as CW; with granules on ven- 
tral surfaces of meri. Pereopod 5 shortest, 
about 1.30 times as long as CW, lacking 
granules on ventral surface of merus. Dac- 
tyl slightly longer than propodus, terminat- 
ing in short corneous claw; with 3 dense 
fringes of setae dorsally, and 1 fringe ven- 
trally. Propodus with dense fringe of setae 
dorsally and ventrally. Carpi of pereopods 
2—4 each with granules and dense fringe of 
setae on dorsal surface; carpus of pereopod 
5 lacking granules, with fringe of setae dor- 
sally. Meri with sparse fringe of setae dor- 
sally. 

Thoracic sternum (Fig. 5a) and abdomen 
with surfaces punctate. Male abdomen (Fig. 
2b, 5b) with short setae marginally on seg- 
ments 1—6 and telson. First segment short, 
narrow. Second segment twice as long as 
first. First and second segments not reach- 
ing coxae of pereopods 5. Third segment 
broadest, overlapping part of coxae of fifth 
pereopods. Fourth and fifth segments sub- 
equal in length. Sixth segment subequal to 
telson in length. Telson bluntly triangular. 

Male pleopod | (Fig. 5c, d) reaching be- 
yond suture separating sternites 4 and 5 
when in situ; medial shoulder with dense 
cluster of spines on ventrolateral face, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


spines diminishing in number, size and 
sharpness distally; distal third with minute, 
broad-based and often blunt spines. Pleo- 
pod 2 (Fig. 5e, f) slightly curved inwardly, 
terminating in subtriangular tip with about 
3 short stiff setae basally on inner angle. 
Gonopores (Fig. 2c) located proximally on 
coxae. 

Female abdomen with first and second 
segments as in male, leaving small portion 
of sternite 8 visible on each side. Gono- 
pores large, transversely ovate. 

Color.—Unknown. 

Etymology.—This species is named in 
recognition of Mrs. Nancy Voss’ late hus- 
band, Dr. Gilbert L. Voss, RSMAS, who 
was Chief Scientist on many of the cruises 
of the R/V Pillsbury. He was also instru- 
mental in making possible the most intense 
and successful deep-sea sampling program 
ever conducted in the tropical Atlantic 
Ocean. The immense number of specimens 
obtained during that program continue to be 
studied, and have considerably enriched our 
knowledge of the fauna from this region of 
the world. 

Distribution.—Caribbean Sea, off Co- 
lombia and Venezuela; Lesser Antilles, off 
southeast coast of Trinidad; and northeast- 
ern coast of South America, off Surinam. 
Depth: 55 to 155 m. 

Similarities.—Given the morphological 
complexities and frequent overlap of char- 
acters among goneplacid taxa, it is useful 
to point out similarities observed between 
Nancyplax vossi, new species, and species 
of other genera in the family. The carapace 
of N. vossi superficially resembles those of 
Carcinoplax species, notably C. barnardi 
Capart, 1951 and C. longispinosa Chen, 
1984. Also, the dense setation of the car- 
pus-propodus-dactyl of N. vossi (Fig. 4) is 
similar to that seen in C. angusta Rathbun, 
1914 (cf. Guinot 1989: 310, fig. 39B). 
However, N. vossi differs from all Carci- 
noplax species in that the latter lack a strid- 
ulatory organ. In addition, the morphology 
of the male pleopods 1 and 2 is quite dif- 
ferent in N. vossi and Carcinoplax species. 


VOLUME 114, NUMBER 4 959 


\ 


\ 
4 
: 
| 


\\\ 
\ 
| 


ae 


\ 


U 


i 


g 
FZ 


yy 


\\ 


CQ QU wv 
Zz 


es 


od 


\ 


\\ 
\ 


Fig. 4. Nancyplax vossi, new species, male 13.9 X 20.2 mm, sta P-752, off Venezuela, USNM 308996. a— 
d, second to fifth right pereopods, lateral view: a, second; b, third; c, fourth; d, fifth. Scale equals 3 mm. 


960 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 5. 
coxae, sternum with abdomen removed leaving right first pleopod. Abbreviations: c, posterior margin of cara- 
pace; cxl-5, coxae of pereopods 1—5; ep4-7, episternites 4 to 7; pl 1, pleopod 1; sl-8, sternites 1—8. Solid black 
indicates membranes. Scales equal 1 mm (a, b), 0.25 mm (c, e), and 0.1 mm (d, f). 


In the new species, pleopod 1 is very slen- 
der distally, and pleopod 2 is short and 
stubby (Fig. 5c—f), whereas in Carcinoplax 
species pleopod | is quite thick distally, and 
pleopod 2 is long and thin (cf. Guinot 
1989). 

Psopheticus species have a stridulatory 
organ very similar to that of N. vossi, but 
otherwise differ markedly. The carapace of 
N. vossi has a long lateral spine (Fig. 1, 2a), 
whereas in Psopheticus species the lateral 


Nancyplax vossi, new species, male 13.9 X 20.2 mm, sta P-752, off Venezuela, USNM 308996. a, 


spine is short (cf. Guinot 1990). Pereopods 
2 to 5 in N. vossi have the meri unarmed, 
and the carpi-propodi-dactyls are densely 
setose (Fig. 4), whereas these pereopods are 
armed with spines and lack dense setation 
in Psopheticus species (cf. Guinot 1990). In 
addition, male pleopods 1 and 2 in N. vossi 
and Psopheticus species differ significantly 
in morphology. 

The male pleopods 1 and 2, abdomen, 
and sternoabdominal structure of Nancy- 


VOLUME 114, NUMBER 4 


plax vossi (Fig. 5c—f) are very similar to 
those of Trizocarcinus dentatus (Rathbun, 
1893) and T. tacitus Chace, 1940 (cf. Guin- 
ot 1969b: 516, figs. 48—51). Male pleopods 
1 and 2 of N. vossi also resemble those of 
Psopheticoides sanguineus Sakai, 1969, the 
only representative of the genus Psopheti- 
coides Sakai, 1969. However, N. vossi dif- 
fers substantially from these species in car- 
apace shape, and armature and proportions 
of pereopods. 

A variety of stridulatory organs have 
been described in the Brachyura (Guinot, 
Dumortier & Dumortier 1960, 1961; Guinot 
1990). The stridulating organ in Nancyplax 
vossi appears to be of the simple kind (cf. 
Guinot 1990) in which the striate plectrum 
on the merus of the cheliped is rubbed 
against the striate pars stridens on the pter- 
igostomial region. 


Acknowledgments 


At RSMAS, we are most thankful to the 
late G. L. Voss for his help and encourage- 
ment for many years, and to his widow 
Nancy Voss, for continuing to encourage 
and support systematic studies of the 
UMML collections. For field work in Co- 
lombia, we thank CIOH for logistic support 
arranged by Navy captains O. Malaver C. 
(director), E. Millan P., and R. Quintero S. 
The Director of INVEMAR, Captain E Ari- 
as I., made possible the use of the B/I An- 
con, and the staff of his institute, in partic- 
ular G. R. Navas, were instrumental in pro- 
cessing the samples. The Smithsonian In- 
stitution supplied financial support for the 
“CIOH-INVEMAR-Smithsonian Expedi- 
tion’ through the office of R. S. Hoffman, 
formerly Smithsonian Assistant Secretary 
for Research. The Office of Fellowships 
and Grants of the Smithsonian Institution 
provided support to RvS in the form of a 
postdoctoral fellowship, and to NHC in the 
form of a short-term visitor travel grant. 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(4):964—971. 2001. 


The systematic position of the genus Basityto Mlikovsky, 1998 (Aves: 
Gruiformes: Gruidae) 


Cécile Mourer-Chauviré 


UMR Paléoenvironnements et Paléobiosphére, Centre des Sciences de la Terre, Université Claude 
Bernard-Lyon 1, 27-43 Boulevard du 11 Novembre, 69622 Villeurbanne Cedex, France. 


Abstract.—The genus Basityto Mlikovsky, 1998 was described as a giant 
Barn Owl and placed in the subfamily Tytoninae, family Strigidae. However 
its morphological characteristics correspond to a crane and it is here transferred 
to the recent genus Balearica, family Gruidae. Other fossil Balearicinae have 
been placed in the genus Probalearica Lambrecht, 1933, type-species Proba- 
learica problematica (Milne-Edwards, 1869). The lectotype of this species, 
here designated, is a partial rostrum which is now known as belonging to 
Palaelodus ambiguus (Phoenicopteriformes: Palaelodidae). The genus Proba- 
learica Lambrecht, 1933 is a junior synonym of Palaeolodus Milne-Edwards, 
1863, and the species Probalearica problematica a junior synonym of Palae- 
lodus ambiguus Milne-Edwards, 1869. The systematic position of the other 
elements attributed to Probalearica problematica, and of the other species at- 
tributed to the genus Probalearica, needs to be revised. Balearica rummeli, 
from the Early Miocene of Germany, is so far the oldest known occurrence of 


the recent genus Balearica. 


In 1998 Mlikovsky described a new ge- 
nus and a new species of Barn Owl, Basi- 
tyto rummeli, in the subfamily Tytoninae 
which he places in the family Strigidae, 
contrary to the most widespread opinion 
(Sibley & Ahlquist 1990). The new species 
was founded on an uncatalogued left hu- 
merus in the private collection of Michael 
Rummel, from the Grafenmiihle 21 locality, 
in Bavaria, Germany, the age of which is 
Early Miocene, Agenian or Orleanian, 
mammal biozone MN 2 or MN 3 (Mein 
1990). I personally examined this holotype, 
which is now catalogued as n° 3/21-0001 in 
Rummel’s collection. A cast of the speci- 
men is deposited in the collection of the 
Université Claude Bernard - Lyon 1, with 
the number FSL 330816. This humerus 
does not belong to a member of the Tyton- 
idae but is very similar to that of the recent 
genus Balearica, in the Gruidae. Anatomi- 
cal terminology follows Baumel & Witmer 
(1993) and, when necessary, Howard 


(1929). Institutional abbreviations: MNHN 
Paris, Muséum national d’ Histoire naturelle 
de Paris; MHN Lyon, Muséum d’ Histoire 
naturelle de Lyon; UCB Lyon, Université 
Claude Bernard - Lyon 1. 


Order Gruiformes 
Family Gruidae 
Genus Balearica Brisson, 1760 
Balearica rummeli (Mlikovsky, 1998), 
new combination 
Figs. 1-3 


Condition of the holotype.—The proxi- 
mal part of the humerus of Basityto has 
been imperfectly restored with plaster on 
the caudal face, so that this part is much 
more proximally straightened that it would 
normally be, although on the cranial face, 
there is a proper contact between the two 
pieces bone. If in the photograph the part 
corresponding to the plaster were removed, 
the true appearance would not be restored 


VOLUME 114, NUMBER 4 


ile 


humeral 
head 


tuberculum 


wwe ventrale 


‘J _— part restored in plaster 


crista 


delto- Be, 


pect. 


Fig. 1. Balearica rummeli, diagram of: the humer- 
us, in dorsal view, showing that the plaster on the cau- 
dal face has given the bone a more straightened aspect. 


because it would still be necessary to rotate 
the proximal part about 21° caudally (Fig. 
1), which would give a rather different as- 
pect. The tuberculum ventrale then seems 
much less proximally elevated in relation to 
the humeral head, and the head forms a 
slight ledge above the incisura capitis and 
the surface of the caudal face. Because of 
this artificially straightened aspect, the hu- 
merus of Basityto does not seem completely 
similar to that in the Gruidae but looks a 
little like that of Xenerodiops mycter (Ras- 
mussen et al. 1987). 

Comparison with Tytonidae and Grui- 
dae, subfamily Balearicinae.—The genus 
Balearica (crowned cranes) shows osteo- 
logical characteristics that are different 
from those of the other genera of the family 
Gruidae, which confirms its allocation to a 
distinct subfamily (Peters 1963, Feduccia & 
Voorhies 1992). Archibald & Meine (1996) 
consider that the genus comprises two spe- 
cies, Balearica pavonina (Linnaeus, 1758) 
and Balearica regulorum (Bennett, 1834) 
although the latter is sometimes considered 
a subspecies of B: pavonina. These two taxa 
have practically the same osteological di- 
mensions (Feduccia & Voorhies 1992). 
Comparison of the humerus of Basityto 
with that of Tytonidae and Balearicinae 


965, 


shows that it is not a barn owl but a 
crowned crane. 

On the caudal face of the proximal part, 
in Basityto, as in Balearica, the caput hu- 
meri is globular and proximally raised, 
while in the Tytonidae it is transversly (..e., 
dorso-ventrally) elongated. In Basityto, as 
in Balearica, the incisura capitis is wide 
and almost oriented along the longitudinal 
axis of the bone, while in the Tytonidae it 
is narrower and obliquely oriented. In the 
Gruidae (Gruinae and Balearica) there is a 
smooth ridge that crosses the incisura cap- 
itis, from the middle of the caput humeri to 
the crus dorsale of the pneumatic fossa. 
This smooth ridge is more globular in the 
Gruinae and more elongate in the Baleari- 
cinae. This ridge is also present in Basityto, 
while it is absent in the Tytonidae. The tu- 
berculum dorsale is very conspicuously 
projecting in Balearica and it is an impor- 
tant characteristic differentiating the Bal- 
earicinae from the Gruinae. It is also strong- 
ly projecting in Basityto, while it is only 
slightly projecting in the Tytonidae. On the 
cranial face, in the Tytonidae, there is a de- 
pression situated between the intumescentia 
humeri and the delto-pectoral crest that is 
proximo-distally elongated. The intumes- 
centia is bordered on its dorsal side by a 
distinct ledge (see Mourer-Chauviré 1987, 
fig. 1). This depression is absent in Balear- 
ica and in Basityto. In Balearica the intu- 
mescentia humeri shows a bulge distally, on 
the ventral side but, this part of the bone is 
missing in Basityto (Figs. 2, 3). 

On the caudal face of the proximal part 
of the shaft, Basityto shows a very elongate 
impression of M. latissimus dorsi in the 
sagittal plane of the bone. This impression 
begins proximally with a tubercle and then 
becomes a thin, elongate impression, that is 
continued distally by a shallow groove. 
Both the tubercle and the muscular impres- 
sion exist in Balearica and in Basityto 
whereas in the Tytonidae the impression of 
M. latissimus dorsi has a completely differ- 
ent shape and is situated on the dorsal side 
of the shaft. The shallow groove is not al- 


966 


ways very pronounced in Balearica, and it 
also exists in the Gruinae (e.g., Grus and 
Anthropoides). 

The distal extremity in the Tytonidae is 
strongly ventrally elongated, whereas it is 
proportionally less elongated in Basityto 
and in Balearica. On the cranial surface, the 
condylus ventralis is globular in Basityto 
and in Balearica, but more elongate in the 
Tytonidae. The ectepicondylar prominence 
is situated more proximally in Basityto and 
in Balearica than in the Tytonidae. The in- 
cisura intercondylaris is deeper in Basityto 
and in Balearica than in the Tytonidae. In 
Basityto, as in the Gruidae, there is a mus- 
cular attachment, situated on the cranial 
face of the epicondylus ventralis, distally to 
the attachment of the anterior articular lig- 
ament. This muscular attachment has the 
shape of a small, deep circular depression. 
In the Tytonidae this muscular attachment 
is situated more ventrally, on the ventral 
face of the epicondylus ventralis (Fig. 3). 
On the caudal surface, in Basityto as in the 
Gruidae, the fossa olecrani is almost invis- 
ible and there are two well marked grooves 
(internal and external tricipital grooves of 
Howard 1929), while in the Tytonidae the 
fossa olecrani is well delimited and there is 
only one visible sulcus, on the ventral side. 

Comparison with recent Gruidae.—Fed- 
uccia & Voorhies (1992) have indicated that 
the humerus of Balearica differs from that 
of Gruinae by the following characteristics: 
humerus with pneumatic fossa less exca- 
vated and with pneumatic foramen smaller; 
crista delto-pectoralis not extended down 
shaft as in Gruinae and intumescentia hu- 
meri relatively less extensive. It can be add- 
ed that the tuberculum dorsale is much 
more proximally projecting, and that the 
distal end is more ventrally elongated in 
Balearica than in Gruinae. 

In Basityto, because of the break, it is not 
possible to see the exact shape of the pneu- 
matic fossa, but it seems that it was shal- 
low. The other characteristics of the Bal- 
earicinae are also found in Basityto. Com- 
pared to Anthropoides virgo, the humerus 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


of which is almost the same size, in Bal- 
earica the crista delto-pectoralis is proxi- 
mo-distally shorter, less extended in the 
cranio-caudal direction, and is inserted 
more on the cranial surface, while in A. vir- 
go it is inserted along the dorso-cranial an- 
gle of the shaft. In these characteristics, B. 
rummeli corresponds perfectly to Balearica. 
In Balearica the condylus dorsalis tapers at 
its proximal end, and sometimes shows a 
slight distalward curvature. This character- 
istic does not appear in Basityto, where the 
end of the condyle seems more rounded, 
but is slightly incomplete (Fig. 3). 

According to these morphological char- 
acteristics, I propose that the genus Basityto 
be placed in synonymy with the genus Bal- 
earica. The species Balearica rummeli dif- 
fers from the recent species B. regulorum 
and B. pavonina by its smaller size. 

Measurements of the humerus (mm), af- 
ter Mlikovsky 1998: estimated total length, 
175; proximal width as preserved, 35; depth 
of head, 10.6; proximal depth from tuber- 
culum ventrale to intumescentia humeri, 
17.2; length from proximal part of caput 
humeri to distal end of crista delto-pecto- 
ralis, 52.5; width and depth of shaft at mid- 
point, 13.1 and 11.2; distal width, ca. 26.5; 
distal depth, 14.6; length from ectepicon- 
dylar prominence to the distal surface of 
condylus dorsalis, 13.0. In B. pavonina the 
humerus length varies from 189.2 to 228.9 
mm (n = 10), and in B. regulorum from 
186.4 to 211.1 mm (nm = 10) (Feduccia & 
Voorhies 1992). The dimensions of B. rum- 
meli are about 90% of the mean dimensions 
of the recent crowned cranes. 

Comparison with fossil Balearicinae.— 
Genus Probalearica Lambrecht, 1933: The 
genus Probalearica was created by Lam- 
brecht (1933) for the species Grus proble- 
matica Milne-Edwards, 1869, described 
from an incomplete os _ premaxillare 
(MNHN, Av 8728 from Saint-Gérand-le- 
Puy (Early Miocene, mammal zone MN 2a) 
and two sterna from Gannat (Late Oligo- 
cene, reference-level MP 30). Cracraft 
(1973) incorrectly considered the anterior 


VOLUME 114, NUMBER 4 967 


Fig. 2. Humeri in caudal view. a, Balearica rummeli (Mlikovsky), left humerus, holotype, Michael Rummel 
collection, n° 3/21.0001. b, Tyto alba, recent, left humerus, Collection UCB, Lyon, n° 245-3. c, Balearica 
pavonina, recent, right humerus, Collection UCB, Lyon, n° I-73. a and c, natural size, b 1.5 X natural size. 


968 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Humeri in cranial view. a, Balearica rummeli (Mlikovsky), left humerus, holotype, Michael Rummel 
collection, n° 3/21.0001. b, Tyto alba, recent, left humerus, Collection UCB, Lyon, n° 245-3. c, Balearica 
pavonina, recent, right humerus, Collection UCB, Lyon, n° I-73. a and c, natural size, b 1.5 X natural size. 


VOLUME 114, NUMBER 4 


part of rostrum to be the holotype, whereas 
it is actually one of three syntypes. I there- 
fore designate the partial rostrum as the lec- 
totype. 

Cheneval & Escuillié (1992) have dem- 
onstrated that this premaxillare does not be- 
long in the Gruidae but is a part of the skull, 
previously unknown, of Palaeolodus am- 
biguus, in the Phoenicopteriformes. The ge- 
nus Probalearica thus becomes a synonym 
of the genus Palaeolodus Milne-Edwards, 
1863, and the species Probalearica prob- 
lematica becomes a synonym of Palaeolo- 
dus ambiguus Milne-Edwards, 1863. 

The sternum illustrated in Milne-Ed- 
wards, 1867-1871, pl. 76, fig. 4, was one 
of the other syntypes of the species ““Grus”’ 
problematica, and is in the collection of the 
Paris MNHN, n° Av 8729, and needs re- 
study. The other syntypical sternum (Milne- 
Edwards, 1867—1871, pl. 76, fig. 3) is miss- 
ing at the present time (Cheneval & Es- 
cuillié 1992). These two sterna were from 
birds the same size as the recent crowned 
cranes, whereas Balearica rummeli is 
slightly smaller. 

Three other species were later placed in 
the genus Probalearica, P. crataegensis, P. 
moldavica, and P. mongolica. The generic 
allocation of these species must be revised. 

““Probalearica”’ crataegensis Brodkorb, 
1963: This species, from the early Miocene 
of Florida is represented only by the distal 
part of a tibiotarsus (Brodkorb 1963). In 
Balearica the tibiotarsus widens medially in 
the distalmost part of the bone, at the level 
of the medial condyle, while in the Gruinae, 
the widening is more progressive and be- 
gins more proximally. In the Gruidae, there 
is a strong tubercle at the distal part of the 
Supratendinal bridge, on its lateral side, 
close to the distal opening of the canalis 
extensorius. In Balearica, on the cranial 
face, the lateral condyle is short and does 
not extend very far proximally, while in the 
Gruinae, it is longer and extends up to the 
level of this tubercle. In Balearica the sul- 
cus extensorius is wide and occupies a large 
part of the cranial face of the distal end, 


969 


while in the Gruidae it is narrower and oc- 
cupies only the medial part of the cranial 
face. In Balearica, the supratendinal bridge 
is wide medio-laterally and extends on the 
lateral side to the tuberositas retinaculi ex- 
tensoris, while in the Gruinae it is much 
narrower in the medio-lateral direction. In 
distal view, in the Gruinae, the condylus 
medialis is more medially projecting than 
in the Balearicinae. In lateral view in the 
Gruinae, the distal outline of the lateral 
condyle forms an indentation that is much 
less pronounced in the Balearicinae. 

The distal part of tibiotarsus of “P.” cra- 
taegensis differs from that of Balearica be- 
cause it does not show a conspicuous wid- 
ening at the level of the condylus medialis, 
the condylus medialis in distal view does 
not show pronounced projecting in the me- 
dial direction, and the indentation of the 
distal outline of the lateral condyle is ab- 
sent. The species may be referable to the 
genus Palaeogrus. 

Aramornis longurio Wetmore, 1926: The 
genus Aramornis, from the earliest middle 
Miocene of Nebraska, was described by 
Wetmore (1926) from the distal part of tar- 
sometatarsus and placed in the family Ar- 
amidae. Olson (1985) has indicated that it 
is a Balearica-like crane and Becker (1987) 
has classified it in the subfamily Balearici- 
nae. There is a large amount of new, un- 
studied crane material, from Oligocene and 
Miocene localities in North America which 
should make it possible to clarify the sys- 
tematic position of “P.” crataegensis and 
A. longurio (Olson 1985, and pers. comm.). 
The size of A. longurio is 67% of the mean 
size of the recent Balearica, and “‘P.”’ cra- 
taegensis is 76%. These two species are 
thus smaller than B. rummeli, the humerus 
of which is 90% smaller than the mean size 
of the recent forms. 

““Probalearica’ moldavica Kurochkin & 
Ganya, 1972: This species, from the Sar- 
matian of Moldavia, was described from an 
incomplete distal portion of tibiotarsus. Its 
age corresponds to the beginning of the late 
Miocene, MN 9 ? (Mlikovsky 1996). This 


970 


specimen does not show much resemblance 
to the Gruidae. The condylus medialis is 
only slightly displaced medially. On the 
cranial face, the condylus lateralis is very 
proximo-distally elongated. The presence of 
a tubercle on the supratendinal bridge is not 
mentioned in the description. The distal 
width is inferior to the distal depth, which 
is not the case in the recent Gruidae. Its size 
is similar to that of the recent Balearica, so 
it is larger than B. rummeli. 

*‘Probalearica’ mongolica Kurochkin, 
1985: This species, from the middle Plio- 
cene of Mongolia, was described from the 
distal part of tibiotarsus, an incomplete dis- 
tal portion of tarsometatarsus, and a quad- 
rate. The tibiotarsus actually shows similar- 
ities with the genus Balearica. On the tar- 
sometatarsus, the three trochleae are bro- 
ken, but it can be seen that trochlea 
metatarsi II is more elongate distally, com- 
pared to trochlea metatarsi III, than in the 
Gruinae, and this characteristic corresponds 
to the Balearicinae. Its size is large, com- 
parable to that of the Common Crane, Grus 
grus, and thus is larger than B. rummeli. 

Balearica exigua Feduccia & Voorhies, 
1992: This species, known from most of the 
skeleton, was described from the early late 
Miocene of Nebraska. It is characterized by 
its small size, which is about 75 to 80% of 
that of the recent Balearica, and thus it is 
smaller than B. rummeli. The humerus dif- 
fers strongly from that of B. rummeli by its 
very robust shaft, by the very strong prox- 
imal projection of the tuberculum dorsale, 
and by the fact that the distal part shows 
almost no ventral widening. 

Comparison with Palaeogrus excelsa 
(Milne-Edwards, 1869).—This species is 
known from the Early Miocene of the 
Saint-Gérand-le-Puy area, MN 2a (Milne- 
Edwards 1867—1871), and is also present in 
the middle Miocene of Sansan, MN 6 (Che- 
neval 2000). It is only mentioned here as a 
matter of interest because it shows a mosaic 
of the morphological characteristics of the 
Balearicinae and the Gruinae. There is a 
large amount of new, unstudied material in 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the collections of MNHN Paris, MHN 
Lyon, and UCB Lyon. The proximal part of 
the humerus, which has not yet been de- 
scribed or illustrated, has the morphological 
characteristics of the Gruinae and differs 
from the Balearicinae. Its size is large, com- 
parable or even slightly superior to that of 
the recent Common Crane, and thus is larg- 
er than B. rummeli. 

Conclusions.—Balearica rummeli, the 
Crowned Crane of Grafenmiihle, Bavaria, is 
as yet the oldest known occurrence of the 
recent genus Balearica, but there is a large 
amount of new, unstudied material, in Eu- 
rope as well as in North America, that may 
provide an earlier appearance. The Balear- 
icinae were widespread in the whole North- 
ern Hemisphere and lived there at least until 
the middle Pliocene of Mongolia, while at 
the present time they only survive in trop- 
ical Africa, south of the Sahara. 


Acknowledgments 


I am grateful to Michael Rummel and 
Ursula Gohlich, from the Institute fiir Pa- 
laontology und historische Geologie, of 
Miinchen, Germany, for giving me the pos- 
sibility to examine the holotype, and to 
Storrs L. Olson, National Museum of Nat- 
ural History, Smithsonian Institution, for 
his comments. The photographs are by Noél 
Podevigne and the drawing by Arlette Ar- 
mand. 


Literature Cited 


Archibald, G. W., & C. D. Meine. 1996. Family Grui- 
dae (Cranes). Pp. 60—89 in J. del Hoyo, A. EI- 
liott, and J. Sargatal, eds., Handbook of the 
Birds of the World, vol. 3. Hoatzin to Auks. 
Barcelona, Lynx Edicions, 752 pp. 

Baumel, J. J., & L. M. Witmer. 1993. Osteologia. Pp. 
45-132 in J. J. Baumel, ed., Handbook of Avian 
Anatomy. Nomina Anatomica Avium, 2nd edi- 
tion. Nuttall Ornithological Club 23, 779 pp. 

Becker, J. J. 1987. Neogene Avian Localities of North 
America. Smithsonian Research Monographs. 
Washington D.C. and London, Smithsonian In- 
stitution Press, 171 pp. 

Brodkorb, P. 1963. Miocene birds from the Hawthorne 


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Foramtion.—Quarterly Journal of the Florida 
Academy of Sciences 26 (2):159—167. 

Cheneval, J. 2000. L-avifaune de Sansan. Pp. 321-388 
in L. Ginsburg, ed., La Faune miocene de San- 
san et son environnement.—Meémoires du Mu- 
séum national d’Histoire naturelle 183, 392 pp. 

, & EF Escuillié. 1992. New data concerning 
Palaelodus ambiguus (Aves: Phoenicopterifor- 
mes: Palaelodidae): Ecological and Evolution- 
ary Interpretations.—Natural History Museum 
of Los Angeles County, Science Series 36:209— 
224. 

Cracraft, J. 1973. Systematics and Evolution of the 
Gruiformes (Class Aves). 3. Phylogeny of the 
Suborder Grues.—Bulletin of the American 
Museum of Natural History 151 (1):1—127. 

Feduccia, A., & M. R. Voorhies. 1992. Crowned 
Cranes (Gruidae: Balearica) in the Miocene of 
Nebraska.—Natural History Museum of Los 
Angeles County, Science Series 36:239—248. 

Howard, H. 1929. The avifauna of Emeryville shell- 
mound.—University of California Publications 
in Zoology 32 (2):301—-394. 

Kurochkin, E. N. 1985. Birds of the Central Asia in 
Pliocene. The joint Soviet-Mongolian paleon- 
tological expedition.—Transactions 26:1—120 
(an Russian). 

, & I. M. Ganya. 1972. Pticy srednego sarmata 
Moldavii (Birds of the middle Sarmatian of 
Moldavia). Pozvonochnye neogena i plejsto- 
cena Moldavii (Vertebrates of the Neogene and 
Pleistocene of Moldavia). Akademia Nauk Mol- 
davskoy SSR:45—70 Gn Russian). 

Lambrecht, K. 1933. Handbuch der Palaeornithologie. 
Berlin, Borntraeger, 1024 pp. 

Mein, P. 1990. Updating of MN Zones. Pp. 73-90 in 
E. H. Lindsay et al., eds., European Neogene 
Mammal Chronology. New York, Plenum Press, 
ix + 658 pp. 


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Milne-Edwards, A., 1867—71. Recherches anatomiques 
et paléontologiques pour servir 4 histoire des 
oiseaux fossiles de la France. Paris, Victor Mas- 
son et Fils. t. 1, 474 pp., atlas pl. 1-96; t. 2, 
627 pp., atlas pl. 97—200. 

Mlikovsky, J. 1996. Tertiary Avian Localities of Mol- 
davia. Pp. 685-691 in J. Mlikovsky, ed., Tertia- 
ry Avian Localities of Europe-—Acta Univer- 
sitatis Carolinae 39 (1995) (3—4):519—848. 

, 1998. A new Barn Owl (Aves: Strigidae) from 
the early Miocene of Germany with comments 
on the fossil history of the Tytoninae.—Journal 
fiir Ornithologie 139:247—261. 

Mourer-Chauviré, C. 1987. Les Strigiformes (Aves) 
des Phosphorites du Quercy (France): Systé- 
matique, Biostratigraphie et Paléobiogéogra- 
phie. Pp 89-135 in C. Mourer-Chauvire, coord., 
L Evolution des Oiseaux d’aprés le Témoignage 
des Fossiles, Table Ronde internationale du 
CNRS.—Documents des Laboratoires de Géo- 
logie de Lyon 99, 248 pp. 

Olson, S. L. 1985. The fossil record of birds. London 
and New York, Academic Press, Avian Biology 
8:79—252. 

Peters, J. L. 1963. Check-list of the birds of the world, 
vol. 2, 2nd edition. Cambridge, Harvard Uni- 
versity Press, 401 pp. 

Rasmussen, D. T., S. L. Olson, & E. L. Simons. 1987. 
Fossil birds from the Oligocene Jebel Qatrani 
Formation, Fayum Province, Egypt.—Smith- 
sonian Contribution to Paleobiology 62: iv + 
20 pp. 

Sibley, C. G., & J. E. Ahlquist. 1990. Phylogeny and 
classification of birds. A study in molecular 
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birds from the Miocene of Nebraska.—Ameri- 
can Museum Novitates 211:1—5. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
114(4):972-987. 2001. 


A new species of tufted-tailed rat, genus Eliurus (Muridae: 
Nesomyinae), from western Madagascar, with notes on the 
distribution of E. myoxinus 


Michael D. Carleton, Steven M. Goodman, and Daniel Rakotondravony 


(MDC) Division of Mammals, Department of Systematic Biology, National Museum of Natural 


History, Smithsonian Institution, Washington, D.C. 20560-0108, U.S.A.; 


(SMG) Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, Chicago, Illinois 


60605, U.S.A. and WWE B.P. 738, Antananarivo (101), Madagascar; 


(DR) Département de Biologie Animale, Université d’ Antananarivo, B.P. 906, Antananarivo 


(101), Madagascar 


Abstract.—A new species of Eliurus, E. antsingy (Muridae: Nesomyinae), 
is described from western Madagascar, only the second one of the genus so far 
known from this vast and biologically underexplored region of the island. The 
distribution of E. myoxinus, the other species known from western Madagascar, 
is also amplified on the basis of new collections that have been made over the 
past decade as interest in small mammals of this region is rekindling. 

Résumé.—Une nouvelle espéce d’Eliurus, E. antsingy (Muridae: Nesomyi- 
nae), est décrite pour la région Ouest de Madagascar. Ce n’est que la deuxieme 
espece du genre connu pour cette vaste région encore biologiquement sous- 
explorée de Madagascar. La distribution de E. myoxinus, | autre espece connue 
pour l’ouest de Madagascar, est aussi étendue grace aux nouvelles collectes qui 
ont été faites au cours de la dernicre décennie car l’intérét porté aux petits 


mammifeéres est rallumé. 


Among Madagascar’s nine endemic ro- 
dent genera, Eliurus, the so-called tufted- 
tailed rats, is the most speciose with nine 
currently recognized species (Carleton 
1994, Carleton & Goodman 1998). Eight of 
these nine species inhabit portions of the 
great humid forest biome (sensu Humbert 
1955) of eastern Madagascar, occurring 
within both lowland and montane rainforest 
associations (for example, Carleton 1994, 
Goodman & Carleton 1996, 1998; Good- 
man et al. 1999). Only one, E. myoxinus, 
the type species of the genus, is known to 
occupy Madagascar’s dryer western and 
southern landscapes, such as dry deciduous 
forest and xerophilous scrub formations 
(Carleton 1994, Goodman et al. 1999). Al- 
though some of the disparity in species 
richness between east and west probably re- 
flects actually lower biodiversity, some may 


be plausibly attributed to the historical in- 
sufficiency of biological survey of small 
mammals in Madagascar’s drier environ- 
ments (see Carleton & Schmidt 1990). For 
example, recent research and collecting in 
western forests have uncovered four new 
species of Microcebus or mouse-lemurs and 
resurrected two others from synonymy 
(Zimmerman et al. 1998, Rasoloarison et al. 
2000). This revision and discovery bring to 
seven the number of Microcebus species 
now recognized from a region that was only 
a few years ago thought to hold one. 

In this paper, we describe a second spe- 
cies of Eliurus from western Madagascar. 
In addition, we report new localities that 
significantly amplify the known geographic 
distribution of E. myoxinus and provide 
western records of two forms apparently re- 
lated to eastern species. 


VOLUME 114, NUMBER 4 


Materials and Methods 


Specimens examined consist principally 
of study skins with associated skulls from 
the following collections, their museum ac- 
ronyms as adopted throughout the report 
contained in parentheses: The Natural His- 
tory Museum (BMNH, formerly British 
Museum of Natural History), London; Field 
Museum of Natural History (FMNH), Chi- 
cago; Muséum National d’ Histoire Naturel- 
le (MNHN), Paris; Département de Biolo- 
gie Animale, Université d’ Antananarivo 
(UA), Antananarivo; National Museum of 
Natural History (USNM, formerly U.S. Na- 
tional Museum), Smithsonian Institution, 
Washington, D.C. Coordinates used to com- 
pose range maps are those given by collec- 
tors for recent collections (see Appendix 1) 
or those estimated by Carleton & Schmidt 
(1990) for older ones. 

A maximum of 7 external and 18 cran- 
iodental variables was recorded in milli- 
meters (mm) for each specimen examined. 
The external variables, most as given by the 
collector, include: total length of body and 
tail (TOTL); head and body length (HBL); 
length of tail vertebrae (TL); hindfoot 
length (HFL, usually without claw for re- 
cent collections); dry hindfoot length 
(DHFL, with claw as measured by Carle- 
ton); ear length (EL); and weight in grams 
_ (WT). The 16 cranial and two dental vari- 
ables (all measured by Carleton) are: 
breadth of the braincase (BBC); breadth 
across both incisive foramina (BIP); 
breadth of the bony palate across the first 
upper molars (BM1s); breadth across the 
occipital condyles (BOC); breadth of the 
rostrum (BR); breadth of the zygomatic 
plate (BZP); depth of the auditory bullae 
(DAB); interorbital breadth (JOB); length 
of bony palate (LBP); length of diastema 
(LD); length of incisive foramen (LIF); cor- 
onal length of maxillary toothrow (LM1-3); 
length of rostrum (LR); occipitonasal length 
(ONL); posterior breadth of the bony palate 
(PPB); postpalatal length (PPL); width of 
the first upper molar (WM1); zygomatic 


973 


breadth (ZB). These were recorded to the 
nearest 0.1 mm, using handheld calipers ac- 
curate to 0.03 mm and following the ana- 
tomical landmarks defined and illustrated in 
Carleton (1994). Morphological terms for 
the muroid skull generally follow Hersh- 
kovitz (1962), Carleton (1980), and Voss 
(1988). 

Due to the small sample sizes encoun- 
tered for most western samples of Eliurus, 
specimens from localites over a general re- 
gion were grouped as operational taxonom- 
ic units (OTUs), as defined below. To better 
gauge the significance of interlocality vari- 
ation among E. myoxinus, four samples of 
a broadly distributed eastern species, E. 
tanala, were included from throughout its 
range. Catalog numbers, full locality data, 
and museum sources of all specimens ana- 
lyzed are provided in Appendix 1. We use 
the following abbreviations for various 
types of Malagasy protected areas: PN, Parc 
National; RE Réserve Forestiere; RNI, Ré- 
serve Naturelle Intégrale; RS, Réserve Spé- 
ciale. 

Eliurus myoxinus.—OTU (1), n = 17, 
from 5 mi E Bevilany, Petriky Forest, and 
Parcel II, PN d’Andohahela. OTU (2), n = 
13, from PN de Isalo, 35 mi E Toliara, Forét 
de Vohibasia, Forét de Vohimena, and Forét 
de Zombitse. OTU (3), n = 9, from Forét 
d’Analabe, Beroboka, Kirindy Forest, Mo- 
rondava, and Tsilambana. OTU (4), n = 12, 
from Forét d’ Analavelona. OTU (5), n = 11, 
from RF de I’ Ankarafantsika. 

Eliurus new species.—OTU (6), n = 2, 
from Bekopaka. OTU (7), n = 3, from the 
RNI de Namoroka. 

Eliurus tanala.—OTU (8), n = 18, from 
PN de Ranomafana and vicinity. OTU (9), n 
= 14, from PN d’Andringitra. OTU (10), n 
= 17, from RS d’ Analamazaotra and vicinity. 
OTU (11), 2 = 6, from RS d’ Anjanaharibe- 
Sud. 

Standard descriptive statistics (mean, 
range, standard deviation) and multivariate 
analyses were calculated based on speci- 
mens of adult age classes (young, full, and 
old) defined on grades of dental wear. Ju- 


974 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 
of Eliurus: left, E. myoxinus (BMNH 47.1603; ONL = 36.7 mm) from 5 mi E Bevilany; right, E. antsingy 
(MNHN 1966.220, holotype; ONL = 41.9 mm) from Bekopaka. (In order to illustrate conformation of a complete 


Dorsal, ventral, and left lateral views (about 1.75) of adult crania and mandibles of western species 


mandible, that of MNHN 1966.2222 was substituted for the partially broken left mandible of the holotype.) 


veniles and subadults, identified by their 
complete or partial gray coats and newly 
erupted or little worn third molars, were ex- 
cluded. Sample statistics for external di- 
mensions were employed in tabular com- 
parisons as a general index of size differ- 
ences among populations and species, but 
were not considered in multivariate sum- 
maries. Derivation of canonical variates and 
principal components was computed using 
only the 18 craniodental measurements, all 
of which had been first transformed to nat- 
ural logarithms. Loadings are expressed as 
Pearson product-moment correlation coef- 
ficients of the principal components or ca- 
nonical variates with logs of the original 
skull variables. All analytical procedures 


were conducted using Systat (version 9.01, 
1999). 


Eliurus antsingy, new species 
Fig. 1, Table 1 


Holotype.—Muséum National Histoire 
Naturelle number 1966.2220, an adult male 
prepared as skin and skull (original number 
86), collected 1 July 1964 by J. M. Klein. 

The round skin is roughly prepared but 
intact, the hind feet unconventionally fold- 
ed forward under the ventrum. It bears two 
tags affixed to the right hind foot, the col- 
lector’s original skin label and a later one 
identifying the collection of the MNHN, 
Paris. The skull is mostly intact, with minor 


VOLUME 114, NUMBER 4 


breakage of the posterior hard palate and 
pterygoid processes; the halves of the lower 
jaw are separated, and both dentaries are 
missing the tips of the coronoid and angular 
processes. The habitat is sparingly noted on 
original tags as “‘en forét.”’ 

No external measurements are found on 
the original labels. Approximate dimen- 
sions, inferred by measurement of the dry 
museum skin, are: total length, 300+ mm; 
tail length, 140+ mm; and hind foot length, 
30+ mm. 

Type locality.—Madagascar, Toliara 
Province, Antsingy Forest, near Bekopaka, 
about 19°07.5'S, 44°49.0’E. 

Original tags read only ““Bekopaka, An- 
tsingy.”’ Tsingy is a type of geological for- 
mation, with a characteristic vegetational 
community, that commences about 3 km to 
the east of the village of Bekopaka and is 
enclosed within a reserve that has been re- 
cently reclassified as the Parc National de 
Bemaraha. Formerly much of the forest in 
this region was within a protected area, var- 
iously called the RNI de Il’ Antsingy or the 
RNI de Bemaraha; only the southern half 
has recently been regazetted as the PN de 
Bemaraha. 

In discussions with M. Dokobe, who col- 
laborated with J. M. Klein during the mis- 
sion to Bemaraha in July 1964, Goodman 
learned that the site where the MNHN spec- 
imens were collected is located to the east 
of the former Bekopaka aviation field (pers. 
comm. in Antsalova, Feb 2001). Dokobe 
confirmed that the animals were trapped in 
typical tsingy forest, near the trail connect- 
ing Bekopaka and Andriandriambe, and we 
estimate the coordinates of the type locality 
to be approximately 19°07.5'S, 44°49.0’E. 

Diagnosis.—A species of Eliurus char- 
acterized by large size (ONL = 40—44 
mm), correspondingly robust molars (LM1- 
3 = 5.5-6.0 mm), relatively long incisive 
foramina (LIF/LD = 56%), and large au- 
ditory bullae. Tail tuft composed of dark 
hairs to the tip, not bicolored. Alveolus of 
lower incisor short, not terminating laterally 
as a capsular process. 


975 


Table 1.—Comparison of selected external and 
craniodental measurements of specimens of Eliurus 
antsingy from the type locality, Bekopaka, and the Ré- 
serve Naturelle Intégrale de Namoroka. 


Bekopaka 
MNHN MNHN RNI de Namoroka 
Variable 1966.2220 1966.2222 (n = 3, 4) 
TL — — 161.7 + 10.2 
150.0-169.0 
HFL — — 30.6 = 0.9 
29.0—31.0 
ONL 41.9 44.1 40.5 = 0.8 
39.8—41.4 
ZB 20.8 21.8 19.1 + 0.4 
18.6-19.4 
BBC 16.3 15.8 15,3) 2 O2 
15.0-15.5 
IOB 5-8) 6.1 6.0 + 0.1 
5.9-6.1 
LR 14.8 15.1 13.4 + 0.6 
12.9-14.0 
BR 7.4 7.5 7.0 = 0.2 
6.7—7.1 
PPL 14.7 15.1 14.5 + 0.2 
14.3-14.7 
LBP 7.8 Voll 7.4 + 0.1 
7.3-7.5 
LIF 6.7 6.8 6.3 + 0.1 
6.2-6.4 
BIF 3.1 2.8 2.6 = 0.1 
2.5—2.7 
LD 11.7 12.4 11.2 = 05 
10.9-11.8 
BMI1s 8.0 8.3 Td) 2 O2 
7.3-7.6 
PPB 6.1 6.0 So) 22 OW 
5.35.7 
DAB 6.0 6.4 5.6 + 0.05 
5.5—5.6 
BZP 3.5 4.3 3.8 + 0.1 
3.7-3.9 
BOC 9.5 9.6 9.0 = 02 
8.89.2 
LM1-3 6.00. 5.65 5.69 + 0.14 
5.51—5.85 
WM1 1.65 1.58 1.49 + 0.03 
1.45—1.53 


Referred specimens.—MNHN 1966. 
2222, an adult female from Bekopaka, also 
noted as from Antsingy; collected 1 July 
1964 by J. M. Klein and prepared as skin 
and skull (original number 85). FMNH 
167563—167566, all young to fully adult fe- 
males from the RNI de Namoroka, collect- 


976 


ed June 1999 by M. Anselme Toto Volahy 
and each prepared as skull with whole car- 
cass in fluid. M. Toto Volahy trapped mam- 
mals in two different sites within the re- 
serve, both in dry deciduous forest. Unfor- 
tunately, the collector’s field tags for the 
specimens listed were lost and it is not pos- 
sible to associate them with the original 
dale 

Distribution.—Known only from the 
type locality and the RNI de Namoroka, 
western Madagascar. 

Description.—Fur texture relatively soft 
and fine. Cover hairs of dorsal pelage bi- 
colored or tricolored: basal three-quarters a 
medium plumbeous gray and the distal one- 
quarter pale buff, some hairs faintly tipped 
with a third band of dusky brown. Cover 
hairs 9-11 mm in length over middle rump. 
Guard hairs medium brown, only slightly 
longer than cover hairs. General effect of 
upperparts a dark drab brown to brownish- 
gray. In specimens from Bekopaka, cover 
hairs of chest, abdomen, and inguinum bi- 
colored, basal half pale gray and distal half 
cream to bright white; hairs over chin and 
throat monocolored white in both speci- 
mens, a tract of creamy white hairs extend- 
ing over the midventer of MNHN 
1966.2222; general effect of underparts a 
medium gray. All specimens from Namo- 
roka with entire venter plain white to pale 
buff, their underparts contrasting sharply 
with dorsum; one individual (FMNH 
167564) with intense buff to pale ochra- 
ceous over thorax, grading to buffy white 
over the abdomen. Tops of forefeet and 
hindfeet covered with silvery white hairs, 
concentrating to form a loose tuft at base 
of claws that extends to or slightly beyond 
their tip. 

Hindfeet comparatively broad, toes ap- 
pearing short proportional to the length of 
the tarsus-metatarsus; HFL, as measured on 
the dry (toes slightly curled) and fluid spec- 
imens, about 29—31 mm, shorter than hind 
foot of E. tanala. Plantar pads six and ar- 
ranged as per genus, but size large, ex- 
tremely bulbous and packed very close to- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


gether; hypothenar pad as big as an inter- 
digital. Pinnae rounded and relatively small 
(about 20 mm as measured on the fluid pre- 
served specimens), in proportion resem- 
bling those of E. majori; color dusky brown 
externally and clothed with fine brown 
hairs, paler brown internally and sparsely 
covered with fine white hairs. One pair each 
of axillary, abdominal, and inguinal mam- 
mae present, total = six (as confirmed on 
MNHN 1966.2222). 

Tail tuft or brush well developed, notice- 
ably evident over distal 40-50% of caudal 
length, hairs over fullest section about 12 
mm long. Tuft dark to tip, composed of pale 
brown to dusky hairs; tuft of holotype with 
some fine white hairs randomly intermixed, 
but not forming a contradistinctive wholly 
white tip as observed in E. tanala. Tail of 
MNHN 1966.2222 and FMNH 167566 ap- 
parently naturally broken at midsection, ter- 
minating as a whorl of dark brown and 
some white hairs. Caudal epidermis medi- 
um gray all around near the base of the tail, 
becoming slightly to heavily mottled, dor- 
sally and ventrally, toward the tip. Scutel- 
lation finely textured. 

Cranium large in size, approximating that 
of E. ellermani or the largest E. tanala, with 
the incisors, zygoma, rostrum and mandible 
correspondingly stout in appearance. Dorsal 
profile of skull appreciably arched, sloping 
gradually toward the rostrum and more 
steeply toward the occiput, not so flat as in 
E. tanala of similar size. Rostrum moder- 
ately long (LR/ONL = 35%), proportions 
typical for the genus; zygomatic arches 
bowed laterally over their midsection, not 
so parallel-sided as in E. myoxinus; brain- 
case relatively wide, rounded and smoothly 
contoured. Incisive foramina bluntly point- 
ed on their anterior and posterior ends, rel- 
atively long and broad for the genus, their 
diastemal expanse (LIF/LD = 56%) in pro- 
portion resembling examples of E. majori 
and E. penicillatus. Posterior palatine fo- 
ramina a single pair of small ovate holes, 
the palatine bones lacking extensive super- 
numerary foramina or slitlike vacuities. 


VOLUME 114, NUMBER 4 


Posterior edge of bony palate about even 
with end of third molars, lacking medial 
spine; anterior rim of mesopterygoid fossa 
nearly straight to gently curved. Alisphe- 
noid strut present on both sides in all spec- 
imens. Subsquamosal fenestra patent but 
small, exposing little or none of the brain 
cavity; hamular process of the squamosal 
short and stout. Auditory bullae absolutely 
the largest in the genus, relatively as large 
as those of E. myoxinus. 

Enamel face of upper and lower incisors 
pale yellow-orange in color. Alveolus of 
lower incisor relatively short, terminating at 
the level of the coronoid process and ap- 
preciably below the sigmoid notch; man- 
dible without prominent capsular process. 
Molar toothrows long (LM1-3 = 5.6—6.0), 
about the size of E. tanala, absolutely and 
relatively shorter than those of E. majori 
and E. penicillatus; upper and lower third 
molars smaller than second. 

Other external, cranial, and dental fea- 
tures as described for the genus (Carleton 
1994, Carleton & Goodman 1998). 

Comparisons.—A combination of large 
physical size, long and wide incisive foram- 
ina, and relatively inflated auditory bullae 
easily distinguishes Eliurus antsingy from 
most members of the genus. For most cra- 
nial variables measured, only individuals of 
E. ellermani and some examples of E. tan- 
- ala attain the same large dimensions. Cra- 
nia of those species, however, possess no- 
ticeably smaller auditory bullae, shorter and 
narrower incisive foramina, a more fenes- 
trated hard palate, and a longer rostrum; 
further, they exhibit a flatter dorsal profile 
compared to the distinctly arched shape of 
E. antsingy. The pinnae and hind feet of E. 
ellermani and E. tanala are also absolutely 
longer. 

Several characteristics of E. antsingy re- 
call examples of E. majori and E. penicil- 
latus. Like those species, its pinnae appear 
small and rounded for the size of the ani- 
mal. Further, the alveolar tract of the lower 
incisor is somewhat short and does not pro- 
duce a conspicuous lateral bulge where it 


977 


terminates within the mandibular ramus; 
and the incisive foramina are comparatively 
long and wide, spanning over 50% of the 
diasternal length. Unlike E. majori and E. 
penicillatus, however, the molar rows of E. 
antsingy resemble most other Eliurus in 
their shorter proportional length, lower 
crown height, and smaller third molars. 

Both E. antsingy and its western conge- 
ner E. myoxinus possess relatively large ec- 
totympanic bullae (large, that is, for the size 
variation observed within Eliurus), but the 
former is not simply an enlarged version of 
the latter (Fig. 1; Tables 1, 3). The cranium 
of E. antsingy lacks the stocky, squarish ap- 
pearance of E. myoxinus, a contrast impart- 
ed by its longer rostrum, comparatively nar- 
row interorbital constriction, and more lat- 
erally bowed zygoma. The two differ in 
configuration of the incisive foramina 
(long, wide in E. antsingy versus short, nar- 
row in E. myoxinus) and development of 
the capsular process (absent in E. antsingy 
versus a moderate bulge in E. myoxinus). 
The pelage of E. antsingy is more drably 
colored than the soft, pale browns of E. 
myoxinus, and its tail tuft is neither as well 
developed in length of individual caudal 
hairs nor in extent of tail vertebrae covered 
as compared with E. myoxinus. 

Visual impressions of differences in skull 
size and shape are generally borne out in 
multivariate analysis of the 18 continuous 
variables recorded for EF. antsingy and rep- 
resentative samples of E. myoxinus and E. 
tanala (Fig. 2, Table 2). The separation of 
E. antsingy from E. myoxinus along the first 
canonical variate principally reflects its 
larger cranial size (correlations mostly large 
and positive); on this axis, the five intact 
specimens of E. antsingy do not stand apart 
from those of FE. tanala. Between-group 
variation summarized by the second canon- 
ical variate involves fewer influential vari- 
ables with only moderate correlations—no- 
tably, inflation of the auditory bullae 
(DAB) and expansiveness of the incisive 
foramina (BIE LIF), and to a lesser extent 
certain indices of cranial breadth (BBC, 


978 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


8 E. antsingy 


E. myoxinus 


CV 2 (14.1 %) 


-8 -6 -4 2 0 2 4 6 8 
CV 1 (64.3 %) 


E. antsingy 6 
7 
1 
2 
E. myoxinus 3 
5 
4 
8 
9 
E. tanala 10 
14 


120 100 80 60 40 20 (0) 
7 2 
Mahalanobis D 


Fig. 2. Results of discriminant function analysis performed on 18 log-transformed craniodental variables, as 
measured on 122 specimens representing 11 OTUs of Eliurus (see Materials and Methods). Top, Projection of 
individual scores onto the first two canonical variables extracted; polygons for E. myoxinus and E. tanala enclose 
the maximal dispersions of specimen scores around a group’s centroid, whereas individual scores are themselves 
plotted for the two samples of E. antsingy. Bottom, Phenogram produced from clustering (UPGMA) of Mahal- 
anobis’ distances among centroids of the 11 OTUs. See Table 2. 


VOLUME 114, NUMBER 4 


Table 2.—Results of discriminant function analysis 
of 11 OTUs representing intact specimens of Eliurus 
myoxinus (n = 50), E. antisingy (n = 5), and E. tanala 
(n = 44) (See Fig. 2). 


Correlations 


Variable CV1 CVv2 
ONL 0.93 0.17 
ZB 0.80 0.19 
BBC 0.49 0.28 
IOB 0.71 0.18 
LR 0.95 0.04 
BR 0.77 0.09 
PPL 0.74 0.20 
LBP 0.91 —0.23 
LIF 0.70 0.44 
BIF 0.34 0.50 
LD 0.93 —0.04 
BMI1s 0.92 0.07 
PPB 0.60 0.33 
DAB 0.06 0.59 
BZP 0.72 0.07 
BOC 0.85 0.32 
LM1-3 0.91 0.14 
WM1 0.87 0.10 
Canonical 

correlation 0.97 0.89 
Eigenvalue 16.5 3.2 


% Variance 64.3 14.1 


BOC, PPB). In each instance, the speci- 
mens of E. antsingy are exceptional for 
their relatively greater development, espe- 
cially as compared with the similarly sized 
E. tanala. 

Remarks.—Acquisition of additional 
specimens of F. antsingy is needed to bring 
taxonomic illumination to the differences 
remarked between the two examples from 
Bekopaka, the type locality, and those from 
the RNI de Namoroka. The latter animals 
are consistently smaller in size (Table 1), 
and their ventral pelage is wholly white to 
pale buff in marked contrast to the gray col- 
ored ventrum of the holotype from Beko- 
paka. The tone of the entirely white venters 
in the Namoroka series resembles the swath 
of white observed on the abdomen of 
MNHN 1966.2222 from Bekopaka, and 
similar variation in ventral pelage colora- 
tion has been noted among populations of 
E. tanala and E. webbi (Carleton 1994, 


979 


Carleton & Goodman 1998). Moreover, 
while acknowledging that existing sample 
sizes of E. antsingy are smaller than desir- 
able, the amount of morphometric diver- 
gence revealed between the Bekopaka and 
Namoroka specimens approximates that 
found among larger geographic samples of 
widely ranging species, such as E. myoxi- 
nus and E. tanala (Fig. 2). In view of their 
other similarities in cranial shape and qual- 
itative features of the skull and skin, we 
provisionally refer the Namoroka speci- 
mens to the new species and urge the need 
for continued study to clarify the interpop- 
ulational variation noted. 

Little is known about the natural history 
of E. antsingy. The two specimens collected 
by Klein near Bekopaka were obtained in 
traps placed on the ground in typical tsingy 
forest habitat (M. Dokobe, pers. comm.). 

The sites within the PN de Bemaraha and 
the RNI de Namoroka are separated by 
about 300 km (by air). The principal ex- 
posed geological formation within the PN 
de Bemaraha is derived from Mesozoic 
limestone, and the forest cover across this 
zone is relatively intact. Thus, we suspect 
that E. antsingy probably occurs throughout 
much of the Bemaraha complex. To the 
north of this zone, a break occurs in the 
Mesozoic limestone formation, which re- 
appears as a series of isolated geological 
islands all the way north to and including 
Namoroka (Du Puy & Moat 1998). Within 
the intermediate zone there are two pro- 
tected areas—the RS de Maningozy, which 
rests on sandstone and unconsolidated 
sands, and the RS de Kasijy, a portion of 
which occurs on Mesozoic limestone. Fur- 
ther survey work is needed to document 
whether FE. antsingy inhabits these inter- 
mediate zones. 

At the type locality of Bekopaka, Klein 
collected a third specimen (MNHN 
1966.2221) of Eliurus together with the two 
examples of E. antsingy described above. 
The fact that the individual was collected at 
a later date (13 September 1964) and on a 
different trip (““Mission no. 7”) suggests 


980 


that Klein captured it in a different area or 
microhabitat. The animal is not represen- 
tative of either E. myoxinus or E. antsingy. 
Overall size is large (ONL = 44.3, LM1-3 
= 5.85), resembling the dimensions of E. 
antsingy and the most robust samples of E. 
tanala (e.g., those from the northern high- 
lands; Carleton & Goodman 1998). Ap- 
proximate external dimensions, as mea- 
sured on the dry skin (no field recorded 
data) are: total length, ca. 340 mm; tail 
length, ca. 160 mm; hind foot length, 34+ 
mm; and ear length, 22+ mm. The upper- 
parts are a pale, grayish-brown; the under- 
parts are entirely creamy white, from the 
chin to the inguinal region, and contrast 
sharply with the brown dorsum. The tail 
tuft is about 70 mm long and consists of 
two colors arranged in three bands: proxi- 
mal (ca. 30 mm) and terminal (ca. 10 mm) 
sections of light brown caudal hairs, sepa- 
rated by a middle segment of purely white 
hairs (ca. 30 mm). Collection of additional 
specimens is needed to determine whether 
this three-banded tuft pattern is typical of 
the population. A short, terminal dark sec- 
tion is observed, albeit rarely, among sam- 
ples of E. tanala, which otherwise possess 
a bicolored tuft composed of dark hairs 
proximally and white hairs towards the tip. 
The pale brown pinnae seem proportionally 
the largest in the genus, particularly in com- 
parison to the specimens of E. antsingy, but 
the wrinkled condition of the dried pinnae 
disallows objective confirmation of our im- 
pression. 

In addition to generous size and relative- 
ly stout construction, the cranium of 
MNHN 1966.2221 exhibits other traits sug- 
gestive of E. tanala or E. ellermani from 
eastern Madagascar. The rostrum is com- 
paratively long with short and narrow in- 
cisive foramina, sharply pointed on both 
ends. Small supernumerary foramina occur 
behind the principal pair of posterior palatal 
foramina, although these are not coalesced 
as elongate vacuities as found in many E. 
tanala. The subsquamosal fenestra is oc- 
cluded, the hamular process thus undefined 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


except at its posteroventral tip where it 
abuts the mastoid chamber. The capsular 
projection of the lower incisor is well de- 
veloped and terminates just below the ven- 
tral rim of the sigmoid notch. Principal 
component analysis of the same craniod- 
ental variables disclosed the same array of 
specimen clusters as obtained in the canon- 
ical variates plot, with MNHN 1966.2221 
positioned at the very periphery of the E. 
tanala constellation (not illustrated; % total 
variance explained on PC I and PC II = 
71.0 and 6.7, respectively). 

Etymology.—The species name is de- 
rived from the Malagasy words ftsingy, 
which refers to the spectacular pinnacle-like 
limestone formation (karst) that occurs to 
the east of Bekopaka (in the Bemaraha re- 
serve complex) and in the RNI de Namo- 
roka, and an, a common prefix which 
means place of. 


Additional Distributional Records of E. 
myoxinus Milne Edwards, 1885 


As understood by Carleton in 1994, the 
geographic range of E. myoxinus was con- 
fined to southern and southwestern Mada- 
gascar, principally along or near the coast 
from the vicinity of Petriky Forest, extreme 
southeastern Madagascar, to the Forét 
d’Analabe, a place just south of the mouth 
of the Tsiribihina River. The few localities 
then known (six) represented dry deciduous 
forest or xerophilous scrub, and none was 
recorded above 245 m. Subsequent reports 
have altered this distributional picture of E. 
myoxinus (e.g., Goodman & Ganzhorn 
1994, Goodman & Rasoloarison 1997, 
Goodman et al. 1999), in particular ampli- 
fying its distribution in the southwestern in- 
terior between the Mangoky and Onilahy 
rivers (Fig. 3). These localities, and others 
listed in Appendix 1, have appreciably 
broadened the known altitudinal span of the 
species in the southwest, now documented 
from near sea level to 1250 m (Forét 
d’Analavelona). In addition, they have re- 
vealed the greater ecological tolerance of 


VOLUME 114, NUMBER 4 981 


Eliurus antsingy A 


Eliurus myoxinus ® 


A 
Tsiribihina 
| 
a 
Mangoky . 


a 


ef” if ‘ 
6 AN: 50 0 200 
: =a seo a 
Onilahy ' WE Kilometers 


Fig. 3. Geographic occurrence of the species Eliurus antsingy and E. myoxinus, the two species so far 
documented as indigenous to western Madagascar. Names refer to the three western rivers discussed in the text, 
and shaded areas indicate highlands above 1200 m. 


982 


the species. Certain habitats above the west- 
ern coastal piedmont and in inland zones of 
moderate elevation (e.g., Vohibasia and An- 
alavelona) are floristically transitional be- 
tween eastern humid forest and western dry 
deciduous forest (Du Puy et al. 1994, Jen- 
kins & Goodman 1999) and suggest accom- 
modation to more mesic conditions than 
previously recorded for the species. 

A recent mission to the summital zone of 
the Analavelona Massif (Nov 2000) uncov- 
ered a remarkable density of Eliurus myox- 
inus, the single nesomyine rodent living in 
pristine forest at 1250 m. During five nights 
of trapping, 500 trap-nights were accrued 
and only two species of rodents were cap- 
tured: 91 individuals of FE. myoxinus and 
116 individuals of the introduced Rattus 
rattus. The abundance of this nesomyine 
species suggests littke or no suppression of 
its population size in the presence of non- 
native Rattus, a finding that contrasts with 
the diminished catch usually recorded for 
native rodents in eastern humid forests 
where Rattus is equally plentiful (Goodman 
1995). Long-term ecological study is need- 
ed to track population fluctuations and de- 
termine whether western Eliurus are less af- 
fected by high densities of Rattus. 

The habitat in the summital zone of An- 
alavelona is mid-altitude montane sclero- 
phyllous forest as defined by Koechlin et 
al. (1974). At this elevation in eastern for- 
est, high levels of small mammal diversity 
have been regularly encountered, with up to 
Six nesomyine species and including as 
many as three sympatric Eliurus (e.g., 
Goodman & Carleton 1998). The Analav- 
elona Forest exhibits floristic and faunistic 
affinities to eastern humid forests (Koechlin 
et al. 1974, Raxworthy & Nussbaum 1997), 
in particular the Central Domain (sensu 
Humbert 1955), and appears to represent a 
mesic relict from a recent geological period 
when the climate over parts of southwestern 
Madagascar was distinctly wetter (Burney 
1993, Goodman & Rakotozafy 1997). On 
the basis of the highly successful capture 
rates of Eliurus at the site, 91 of 207 ro- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


dents trapped and all referable to E. myox- 
inus, the sympatric occurrence of a conge- 
ner seems unlikely. In spite of the biogeo- 
graphic affinities demonstrated between the 
Analavelona Forest and eastern humid for- 
ests for certain organisms, such past links 
cannot be conjectured based on the rodent 
community as presently understood. How- 
ever, evidence for past east-west biotic con- 
nections is found in the nearby PN de Isalo, 
where a form related to eastern E. majori 
has been identified (Jansa & al. 1999). 

We here report additional geographic oc- 
currences of E. myoxinus that considerably 
extend its range northwards to the region of 
the RF de Jl Ankarafantsika (16°20.3’S), 
Mahajanga Province, a distance of about 
480 km from the species’ previous north- 
ernmost locality (Forét d’Analabe, 
19°58’S). Specimens of E. myoxinus were 
first collected in January 1991 (Ampijoroa, 
70 m) and later in February 1997 (5 km 
SSW Ampijoroa, 160 m). The natural 
wooded habitat in this reserve consists of 
dry deciduous forest, and the site of the 
1997 collections was distinctly degraded 
due to human activities. Within the reserve 
and its vicinity, E. myoxinus occurs in sym- 
patry with Macrotarsomys bastardi and M. 
ingens. 

The northern samples of E. myoxinus av- 
erage slightly but consistently smaller in 
most dimensions of the skull compared 
with southern populations (Table 3). Still in 
aspects of cranial shape, the series from An- 
karafantsika exhibits the blocky construc- 
tion characteristic of E. myoxinus, such as 
the parallel-sided zygoma, rectangular 
braincase, and relatively wide interorbital 
region. The dorsal pelage, however, is gen- 
erally more saturated and darker in shade 
than the sandy brown typical of E. myoxi- 
nus, though the texture is similarly fine and 
the length relatively short. The tail tuft is 
long and bushy, as is characteristic of the 
species, but the caudal hairs are not quite 
so long, perhaps a reflection only of its 
smaller body size. The pelage of one animal 
(UADBA 9936) in the series is inordinately 


VOLUME 114, NUMBER 4 


Table 3.—Comparison of selected external and 
craniodental measurements for the holotype (MNHN 
1886.1120) and two population samples of Eliurus 
myoxinus (sample statistics include the mean + 1 SD 


and the observed range). 


MNHN Analavelona Ankarafantsika 
Variable 1886.1120 (n = 12) (n = 8-11) 
HBL — 127.7 + 6.4 123.6 + 8.7 
117-136 109-130 
TL —_— 145.5 + 12.8 IB27 Ss @,5) 
125.0-167.0 115-142 
HFL — 25.4 + 0.8 M3 == OS) 
24.0—26.0 23.0—24.0 
EL — 23.7 = 0.8 20.0 + 1.9 
23.0—25.0 18.0—23.0 
WT — 65.5 + 8.3 SOM eal 
51-75 36-79 
ONL 34.2 86:1 22 12 34.5 + 0.8 
34.0-37.8 33.0-35.3 
ZB 17.6 17.8 + 0.8 17.6 + 0.4 
16.7—19.0 17.1—18.2 
BBC 13.6 14.0 + 0.4 1357 25 02 
13.1—14.5 13.5-14.0 
IOB 5.5 5.6 + 0.1 5.4 + 0.1 
5.2—5.7 5.35.6 
LR 11.4 12.4 = 0.7 11.4 + 0.2 
10.7-13.3 11.2-11.7 
BR 6.6 7 = 02 6.4 + 0.3 
6.3—7.0 6.1—7.0 
PPL — I 22 7 12.7 + 0.4 
11.8-14.0 12.0-13.1 
LBP 7.1 7.6 = 0.5 7.0 = 0.3 
6.7-8.3 6.6—7.7 
LIF 5.4 5.0 + 0.3 49 + 0.2 
4.5-5.7 4.6-5.1 
BIF 2.0 Dae (al 3) 22 (0), 
1.9-2.5 2.2—2.4 
LD 10.1 10.3 + 0.6 10.0 = 0.3 
9.1-11.0 9.5-10.4 
BMIs 6.7 Foil = O38 6.6 + 0.2 
6.5—7.5 6.4-6.9 
DAB 5.0 So) 2= (03) 5.1 = 0.1 
5.2-5.9 5.0-5.3 
BZP 3.1 3.3 + 0.1 3.4 + 0.2 
3.0-3.6 3.1-3.7 
BOC — 7.9 + 0.3 Vl = OB 
7.3-8.3 7.1-8.1 
LM1-3 4.90 SII) 23 (075) 4.72 + 0.10 
4.74—5.55 4.57-4.90 
WM1 1.27 1.41 + 0.07 1.28 + 0.07 
1.30-1.52 1.18-1.38 


983 


“bleached,” its dorsum appearing uniform- 
ly tan or light tawny. 

Although the Ankarafantsika series is 
nearer the small end of size variation within 
the species, multivariate analyses empha- 
size the basic craniodental conformity of 
the sample to others of FE. myoxinus (Fig. 
2). The amount of inter-sample differentia- 
tion among all OTUs of E. myoxinus, in- 
cluding that from Ankarafantsika, resem- 
bles the pattern of variation shown by sam- 
ples of a broadly distributed eastern species, 
E. tanala. Recent molecular work on ne- 
somyine rodents has included samples 
across the range of E. myoxinus (including, 
from north to south, Ankarafantsika, Kir- 
indy, Isalo, Zombitse, and Andohahela [par- 
cel II]—see Appendix 1), and the amount of 
sequence divergence and resulting clado- 
gram are consistent with the hypothesis that 
these various populations represent a single 
species (Jansa 1998, Jansa & al. 1999). 

Two other noteworthy examples of E. 
myoxinus have been recently (Nov-Dec 
1999) captured at a second northern locale, 
the Forét d’ Ankazotsihitafototra (1150 m) 
in the RS d’Ambohyanahary. This site 
(18°15.7’'S) lies geographically intermediate 
to Forét d’Analabe and the RF de 
1’ Ankarafantsika and suggests that future 
biological survey will disclose a broad dis- 
tribution of the species throughout western 
landscapes. The Forét d’ Ankazotsihita- 
fototra is situated at the very western edge 
of the Central Highlands, lying to the north 
and east of the Bemaraha reserve complex, 
and consists of humid forest of the Central 
Domain (sensu Humbert 1955). Faunisti- 
cally, the Forét d’Ankazotsihitafototra is a 
mixture of eastern and western elements. 
Within the RS d’ Ambohijanahary, EF. myox- 
inus was taken sympatrically with two in- 
dividuals (FMNH 167546, 167555) of an E. 
majori-like form. They are appreciably 
smaller than typical EF. majori with stronger 
contrast of the dorsal-ventral pelage color. 
While further study is required to properly 
assess the status and relationship of this 
form, its trenchant features recall species 


984 


(E. majori and E. penicillatus) that were 
previously known only from eastern Mad- 
agascar. 


Discussion 


The unique combination of morphologi- 
cal features exhibited by E. antsingy pro- 
vides insufficient basis for meaningful com- 
ment on the new species’ closest relatives. 
We do suspect that, although presumably 
western in its distributional setting like E. 
myoxinus, E. antsingy is not specially re- 
lated to this more widely known western 
form. Firmer understanding of their level of 
relationship within the radiation of Eliurus 
must await the acquisition of additional 
specimens, recourse to other sources of tax- 
onomic information, and broader intrage- 
neric comparisons. Other geographic par- 
allels are known within Madagascar’s small 
mammal fauna. For example, molecular ev- 
idence suggests that the species of mouse 
lemur known from the Bemaraha area, Mi- 
crocebus myoxinus, is one of the sister taxa 
to the eastern M. rufus group rather than the 
widespread M. murinus group (Yoder et al. 
2000). 

As currently understood, the taxonomic 
diversity of Madagascar’s indigenous ro- 
dents is concentrated along the eastern one- 
third of the island as compared to its spa- 
cious western landscapes (Table 4). Seven 
genera representing seventeen species have 
been documented to date in the east versus 
four genera representing eight species in the 
west. Such a contrast in species richness 
could be plausibly attributed to the circum- 
stantial interplay of the island’s physical ge- 
ology and regional climatic patterns, and at- 
tendant opportunities for the stratification of 
plant communities, greater ecological com- 
plexity, and geographic isolation that fol- 
low. 

However, while recent biological inven- 
tories have substantially bolstered our 
knowledge of nesomyine distributions in 
eastern Madagascar (Goodman & Carleton 
1996, 1998, Goodman et al. 1996, 1999, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 4.—Tabulation of nesomyine species known 
from eastern versus western landscapes of Madagascar. 


Eastern Westem 


Brachytarsomys albicauda 
B. villosa** 

Brachyuromys betsileoensis 
B. ramirohitra 

Eliurus ellermani* 


Eliurus antsingy* 

E. aff. majori 

E. myoxinus 

E. aff. tanala 
Hypogeomys antimena 
Macrotarsomys bastardi 
M. ingens 

Nesomys lambertoni** 


. grandidieri* 

. majori** 

. minor 

. penicillatus** 

. petteri* 

. tanala** 

webbi** 
Gymnuromys roberti 
Monticolomys* koopmani* 
Nesomys audeberti** 

N. rufus 

Voalavo* gymnocaudus* 


Rata ase 


* = Described as new since 1994. 
**« = Resurrected from junior synonymy since 1994. 


Carleton & Goodman 2000), vouchered 
documentation of nesomyine occurrences in 
the west has only started to fill in the sig- 
nificant geographic gaps identified by 
Carleton & Schmidt (1990). Until now, Eli- 
urus myoxinus, described by Milne Ed- 
wards in 1885, has remained the only spe- 
cies of that genus recorded from western 
environments; over just the past six years, 
three eastern species of Eliurus have been 
newly described and another four recently 
resurrected from junior synonymy as valid 
(Carleton 1994, Carleton & Goodman 
1998). The diagnosis of E. antsingy, re- 
ported herein, and the presence of autoch- 
thonous genera like Hypogeomys and Ma- 
crotarsomys, together suggest the rich po- 
tential for future discoveries among the is- 
land’s western habitats. The recovery of 
additional species from Ambohijanahary 
and Bekopaka that appear to have affinity 
to forms of eastern Eliurus, like the recent 
report of an E. majori-like form from Isalo 
(Jansa & al. 1999), further attest our im- 
perfect knowledge of nesomyine distribu- 
tions in western Madagascar. In view of the 
many regions that still lack thorough site 


VOLUME 114, NUMBER 4 


survey, any meaningful conclusion on the 
degree of east-west biodiversity contrasts 
must await this firmer foundation of field 
inventory and systematic study. 


Acknowledgments 


We especially thank Michel Tranier, Mu- 
séum National d’Histoire Naturelle, Paris, 
and Paula Jenkins, The Natural History 
Museum, London, for allowing us to ex- 
amine specimens under this care. John 
Phelps and William Stanley, Field Museum 
of Natural History, promptly and efficiently 
handled loan transactions to the senior au- 
thor, and Dave Schmidt (USNM) undertook 
the cranial photography and distributional 
map. M. Dokobe of Antsalova kindly pro- 
vided information on the sites that J. M. 
Klein visited in July 1964. Funding for re- 
cent field work in western Madagascar was 
provided by the World Wide Fund for Na- 
ture, The National Geographic Society 
(6338-98 and 6637-99), and the John D. 
and Catherine T. MacArthur Foundation. 
Finally, we appreciate the careful reviews 
and helpful suggestions offered by Sharon 
Jansa and Paula Jenkins. 


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Appendix 1 


Listed below are specimens of Eliurus that formed 
the basis for the morphological comparisons, sample 
statistics, morphometric analyses, and mapped distri- 
butions presented above. 

Eliurus myoxinus.—Fianarantsoa Province: Parc 
National de Isalo, 3.8 km NW Ranohira, along Na- 
maza River, 800 m; 22°32.4’S, 45°22.8'E (FMNH 
166079—166081). Mahajanga Province: Forest Station, 
Ampijoroa, 70 m, 16°15’S, 46°48’E (DR 165; USNM 
576754—576756); Réserve Forestiére de 1 Ankara- 
fantsika, 5 km SSW Ampijoroa, 160 m, 16°20.3’S, 
46°47.6'E (UADBA 9935-9943); Réserve Spéciale 
d’ Ambohijanahary, Forét d’ Ankazotsihitafototra, 1150 
m; 18°15.7’S, 45°25.2’E (FMNH 167545, 167547). To- 
liara Province: Forét d’Analabe, 60 km N Morondava 
(MNHN 1980.290, 1982.988); Forét d’ Analavelona, 
12 km NW Andranoheza, 1050 m, 22°40.7’S, 
44°11.5’E (FMNH 161578—-161589); Réserve Naturel- 
le Intégrale d’ Andohahela, parcel I, 7.5 km ENE Ha- 
zofotsy, 120 m, 24°49.0'S, 46°36.6'E (FMNH 
156630); Beroboka, 40 mi N Morondava (BMNH 
47.1608, 47.1609, 1987.50); 5 mi E Bevilany, Amboy- 
ombe-Fort Dauphin road, the hills, 800 ft (BMNH 
47.1600—47.1607, 66.2746); Kirindy Forest, 60 km 
NE Morondava, 20°04'S, 44°49'E (FMNH 154632, 
154633); Lamboromakandro, Forét de Sakaraha 
(MNHN 1961.220); Morondava (MNHN 1973.516); 
Petriky Forest, 5-7 km SE Manambaro, 20 m, 25°04’S, 
46°53'E (USNM 578679-578687); Tsilambana 
(MNHN 1886.1120, holotype); 35 mi E Tulear 
(BMNH 47.1610, 47.1611, 47.1611a); Forét de Vohi- 
basia, 59 km NE Sakaraha, 780 m, 22°27.5’S, 
44°50.5'E (FMNH 156183, 156191); Forét de Vohi- 
mena, 35 km SE Sakaraha (FMNH 156192—156194); 
Forét de Zombitsy, 870 m, 22°51'S, 44°43'E (FMNH 
151951, 151952). 

Eliurus tanala.—Antsiranana Province: Réserve 


VOLUME 114, NUMBER 4 


Spéciale d’ Anjanaharibe-Sud, 6.5 km SSW Befingitra, 
875 m, 14°45.3'S, 49°30.3'E (FMNH 154049); Ré- 
serve Spéciale d’ Anjanaharibe-Sud, 9.2 km WSW Be- 
fingitra, 1260 m, 14°44.7'S, 49°27.7'E (FMNH 
154050, 154051, 154249, 154250; UA-SMG 6979). 
Fianarantsoa Province: Ambodiamontana, 7 km W of 
Ranomafana, 950 m, 21°16’S, 47°26’E (USNM 
448981—448990, 448997, 449250); 3 km NNW Voh- 
iparara, 1225 m, 21°13’S, 47°22’E (USNM 449251— 
449255): 1 km NW Andrambovato, 875 m, 21°30’S, 
47°25'E (USNM 449256); Réserve Naturelle Intégrale 
d’ Andringitra, 38 km S Ambalavao, ridge east of Vol- 
otsangana River, 1625 m, 22°11’S, 46°58’'E (FMNH 


987 


151691, 151692, 151744, 151880, 151881); Réserve 
Naturelle Intégrale d’Andringitra, 40 km S Ambala- 
vao, along Volotsangana River, 1210 m, 22°13’S, 
46°58’E (FMNH 151690, 151743, 151873, 151874, 
151897); Réserve Naturelle Intégrale d’ Andringitra, 43 
km S Ambalavao, junction of Sahanivoraky and Sa- 
havatoy rivers, 810 m, 22°13’S, 47°00’E (FMNH 
151687, 151688, 151869, 151870). Toamasina Prov- 
ince: 10 mi NW Lohariandava, 1500 ft (BMNH 
47.1573); Périnet, near Moramanga, 3000 ft (BMNH 
47.1557, 47.1560—47.1562, 47.1564—47.1568, 
47.1571, 47.1572; MNHN 1961.176); 1 km E Périnet 
(USNM 341826); 2 km E Périnet (USNM 328828, 
328829): 13 km E Périnet (USNM 341827). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


114(4):988—992. 2001. 


The lophialetid ceratomorph Eoletes (Mammalia) from the Eocene of 


the Zaysan Basin, Kazakstan 


Spencer G. Lucas, Luke T. Holbrook, and Robert J. Emry 


(SGL) New Mexico Museum of Natural History, 1801 Mountain Road N.W., Albuquerque, 


New Mexico 87104, U.S.A; 


(LTH) Department of Biological Sciences, Rowan University, 201 Mullica Hill Rd., Glassboro, 


New Jersey 08028-1701, U.S.A.; 


(RJE) Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, 


Washington, D. C. 20560-0121, U.S.A. 


Abstract.—The “hyrachyid’’ ceratomorph Subhyrachyus tshakpaktasensis 
Gabuniya, was named for a nearly complete skull from the Zaysan Basin of 
eastern Kazakstan. This skull is from the Eocene (Arshantan) Chakpaktas For- 
mation at the Mozhevelnik locality on the Kalmakpay River. Re-examination 
of the skull indicates it belongs to the lophialetid genus Eoletes Biryukoy, 
based on the following diagnostic features: shallow nasal incision only retracted 
to above the canine, long and shallow maxillary fossa, anterior edge of orbit 
above M2, no modifications of rostrum as in Lophialetes, short postcanine 
diastema, complete u-shaped lophoid loops on P3-4, and relatively lophodont 
cheek teeth. The features that distinguish E. tshakpaktasensis from E. gracilis 
Biryukov, (type species of Eoletes) and Chinese FE. xianensis Zhang & Qi, are 
minor (upper cheek teeth relatively wider and more prominent premolar ecto- 
loph ribs in E. tshakpaktasensis), but we retain all species as valid pending a 
larger sample with which to document better dental variability in Eoletes. E. 
tshakpaktasensis is the first record of Eoletes from the Zaysan Basin and ex- 
tends the temporal range of the genus from the Irdinmanhan back to the Ar- 


shantan. 


The Zaysan Basin of eastern Kazakstan 
(Fig. 1) yields an extensive fossil record of 
Eocene mammals that represent the Arshan- 
tan, Irdinmanhan, and Ergilian land-mam- 
mal “‘ages” (lma) (e.g., Russell & Zhai 
1987, Lucas et al. 2000, Lucas 2001). New 
discoveries in the Zaysan Basin continue to 
augment our knowledge of Asian Eocene 
mammals. Gabuniya (1999) described a new 
perissodactyl genus and species, Subhyrach- 
yus tshakpaktasensis, from the oldest Eocene 
strata known in the Zaysan Basin. As the 
name indicates, Gabuniya considered this 
new taxon to be a member of the cerato- 
morph family Hyrachyidae and concluded 
that it was part of an Asian diversification 
of hyrachyids distinct from the well-docu- 


mented North American forms, discussed by 
Wood (1934) and Radinsky (1965, 1967). 
However, reexamination of the holotype and 
only specimen of Subhyrachyus tshakpak- 
tasensis indicates that it should be reas- 
signed to the lophialetid genus Eoletes. Here 
we document this reassignment and briefly 
discuss its implications. 

Abbreviations used.—When used in den- 
tal notations, uppercase letters denote upper 
(premaxillary and maxillary) teeth, and 
lowercase letters denote lower (dentary) 
teeth. Institutional abbreviations are: 
IPGAN = Institute of Paleobiology, Geor- 
gian Academy of Sciences, Tbilisi; KAN = 
Institute of Zoology, Academy of Sciences 
of the Republic of Kazakstan, Almaty. 


VOLUME 114, NUMBER 4 


Miles 0 100 200 300 
L mn = -L 1 


i j 
Kilometers 0 100 200 300 400 500 


Russia 


ra 
=e, 
S 
Ss 
. "sy" 
Sea 
Aktyubinsk 


989 


“wv 


Pe: ~'e ) 
vn. > 7Petropaviovsk’. wv. 


Neos 


@ 
Semipalatinsk 


Mongolia ¥ 


Kazakstan 


Kyzyl-Orda 


Necsae 
fee S) 

els ( < ‘Uzbekistan ¢ 
: % 


Turkmenistan iN ; 
ao =. a we . 
SS 


Tashkent“{- 
fy ENTS a 


Alma-Ata 2 ie 
wa 
Kyrgyzstan 
el ; 
TS ENS 


~~ 


China 


A 
Ls a 


. Nw AS 5 
i Afghanistan S SSrakitan 


Fig. 1. 


Systematic Paleontology 


Family Lophialetidae Matthew & Granger, 
E25 
Genus Eoletes Biryukov, 1974 
Eoletes tshakpaktasensis (Gabuniya, 1999) 
Figs. 2-3 


Subhyrachyus tshakpaktasensis Gabuniya, 
1999:561, figs. 1-3. 


Holotype.—IPGAN Z402, nearly com- 
plete skull with right P3-M3 and left P2- 
M3 (Gabuniya, 1999, figs. 1-3) (Figs. 2-3). 

Type locality and horizon.—Mozhevel- 


Map of Kazakstan showing location of Zaysan Basin. 


nik locality on the Kalmakpay River, Zay- 
san Basin, Kazakstan (UTM Zone 45, 
383463E, 5256668N, datum WGS84). 

Referred specimen.—Only known from 
the holotype. 

Description.—Gabuniya (1999) provided 
a detailed and accurate description of 
IPGAN Z402, obviating the need for exten- 
sive description here. Instead, we list the 
salient features critical to determining the 
systematic position of this fossil below. 

The nasal incision is very shallow, only 
retracted to above the canine: 


990 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. 
tos), and left lateral (C; stereophotos) views. 


The facial portion of the maxilla bears a 
moderately deep preorbital fossa: 

The nasals extend far anteriorly and ap- 
parently contact the premaxilla: 

The supraorbital processes are damaged, 
so that their complete extent cannot be de- 
termined: 

The P3 and P4 each bear a protoloph and 
metaloph that are joined lingually, forming 
a U-shaped loop. This is the “‘complete lo- 
phoid loop” character of Hooker (1989, p. 
88, table 6.1, character 21): 

The upper cheek teeth are relatively 
lophodont, with prominent, sharp ecto- 
lophs, prominent molar parastyles, lingually 
deflected molar metacones, and sharp, well- 
developed molar protolophs and metalophs 


IPGAN Z402, skull of Eoletes tschakpaktasensis in dorsal (A; stereophotos), ventral (B; stereopho- 


connecting the lingual cusps to the ecto- 
loph: 

Measurements of the cheek teeth of 
IPGAN Z402 Gin mm) are: P2 L = 8.8, W 
= 8.6; P3 L = 9.4, W = 10.4; P4 L = 95, 
W = 11.4; M1 L = 12.3, W = 14.1; M2 
L = 14.6, W = 15.5; M3 L = 13.9, W = 
14.7: 


Discussion—The U-shaped lophs of the 
premolars are a synapomorphy of the Lo- 
phialetidae (Hooker 1989, Lucas et al. 1997) 
and indicate that IPGAN Z402 is a lophiale- 
tid. In hyrachyids, the premolar metalophs 
are short and do not contact the protoloph. 
Lophialetids include the genera Lophialetes, 
Schlosseria, Breviodon, and Eoletes. Lophi- 


VOLUME 114, NUMBER 4 


Fig. 3. 


991 


1 cm 


Occlusal views of upper cheek teeth of Eoletes. A, IPGAN Z402, Eoletes tshakpaktasensis, right P2- 


M3. B) KAN 5088/69, Eoletes gracilis, left P1-M3 (photo reversed). 


aletes differs from the others in possessing 
a narial incision that is retracted far poste- 
riorly, similar to what is seen in living Tap- 
irus. IPGAN ZA02 is similar to many prim- 
itive perissodactyls, including Hyrachyus 
and lophialetids other than Lophialetes and 
Schlosseria, 1n possessing an unretracted 
narial incision. Among lophialetids without 
a retracted narial incision, only Eoletes pos- 
_ sesses a well-developed maxillary fossa (Lu- 
cas et al. 1997). Eoletes also possesses a 
long supraorbital process. 

The morphology of the premolars indi- 
cates that IPGAN Z402 is a lophialetid, not 
a hyrachyid. The absence of a retracted nar- 
ial incision indicates that this specimen 
does not belong to Lophialetes or Schlos- 
seria. IPGAN ZAO02 is very similar in its 
cranial morphology to specimens of Eoletes 
gracilis from the Shinzaly locality in east- 
erm Kazakstan, particularly in terms of the 
morphology of the preorbital fossa and den- 
tition (Biryukov 1974, Reshetov 1979, Lu- 
cas et al. 1997, figs. 3—4). Unfortunately, 
the poor preservation of the supraorbital 
processes prevents any assessment of 


whether IPGAN Z402 is similar to the 
Shinzaly specimens of Eoletes in this fea- 
ture. Nevertheless, we feel that the similar- 
ities between these specimens justify as- 
signing IPGAN Z402 to Eoletes. 

IPGAN Z402 does differ from described 
species of Eoletes (E. gracilis and E. xia- 
nensis Zhang & Qi, 1981) in some minor 
features. Thus, IPGAN Z4072 has relatively 
wider cheek teeth, more prominent ectoloph 
ribs on the upper premolars, and relatively 
larger upper molar parastyles than other 
specimens of Eoletes. Because little is 
known of dental variation in Eoletes, it is 
difficult to evaluate the taxonomic signifi- 
cance of these differences. Although we 
suspect that these differences may be found 
in the future to fall within the range of in- 
traspecific variation for this taxon, we take 
the conservative course and tentatively re- 
tain Gabuniya’s species, and we refer 
IPGAN ZA402 to Eoletes tshakpaktasensis. 


Biostratigraphy 


The holotype of E. tshakpaktasensis is 
the first record of Eoletes from the Zaysan 


992 


Basin and is the oldest record of the genus. 
It is derived from the Chakpaktas svita, 
which is of Arshantan age (Lucas 2001). 
The type locality and only record of E. 
gracilis, at Shinzaly, is younger, of Irdin- 
manhan age (Lucas et al. 1997, Lucas 
2001). The only occurrence of E. xianensis 
in the Bailuyuan Formation of Shaanxi is 
Irdinmanhan or possibly slightly younger 
(Lucas et al. 1997). Eoletes thus remains a 
rare tapiroid from the Eocene (Arshantan- 
Irdinmanhan) of Asia. 


Acknowledgments 


The National Geographic Society and the 
Charles D. Walcott Fund of the Smithson- 
ian Institution supported this research. Le- 
onid Gabuniya generously allowed us to 
study the holotype of ““Subhyrachyus”’ 
tshakpaktasensis. We thank P. Holroyd and 
an anonymous reviewer for comments that 
improved the manuscript. 


Literature Cited 


Biryukoyv, M. D. 1974. Novye rod semeystva Lophi- 
aletidae iz eotsena Kazakhstana [New genus of 
the family Lophialetidae from the Eocene of 
Kazakhstan].—Akademiya Nauk Kazakhskoy 
SSR Institut Zoologii Materiali po istorii fauni 
I flori Kazakhstana [Academy of Sciences of the 
Kazakh SSR Institute of Zoology Material for 
the History of the Fauna and Flora of Kazakhs- 
tan] 6:57—73. 

Gabuniya, L. K. 1999. On a new member of the Hy- 
rachyidae (Perissodactyla, Ceratomorpha) from 
the Eocene of the Zaisan depression (eastern 
Kazakstan).—Paleontological Journal 33:561— 
565. 

Hooker, J. J. 1989. Character polarities in early peris- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


sodactyls and their significance for Hyracoth- 
erium and infraordinal relationships. Pp. 79— 
101 in D. R. Prothero & R. M. Schoch, eds., 
The evolution of perissodactyls. Oxford Uni- 
versity Press, 537 pp. 

Lucas, S. G. 2001. Gobiatherium (Mammalia: Dino- 
cerata) from the middle Eocene of Asia: Tax- 
onomy and biochronological significance —Pa- 
laontologische Zeitschrift 74(4):591—600. 

, R. J. Emry, & B. U. Bayshashov. 1997. Eo- 

cene Perissodactyla from the Shinzaly River, 

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leontology 17:235—246. 

, V. Chkhikvadze, B. Bayshashovy, L. 
A Taleo P. A. Tleuberdina, & A. Zhaman- 
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desch & K.R. Kelts, eds., Lake Basins Through 
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Matthew, W. D., & W. Granger. 1925. The smaller pe- 
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Radinsky, L. B. 1965. Early Tertiary Tapiroidea of 
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. 1967. Hyrachyus, Chasmotherium, and the 
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Reshetov, V. J. 1979. Rannetretichnyie tapiroobrazniye 
Mongolii i SSSR [Early Tertiary tapiroids of 
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golskaya Paleontologicheskaya Ekspeditsiya 
11:1-144. 

Russell, D. E. & R. Zhai. 1987. The Paleogene of Asia: 
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Asiatica 19:214—217. 


PROCEEDINGS 


of the 


Biological Society of 


Washington 


VOLUME 114 
2001 


Vol. 114(1) published 19 April 2001 Vol. 114(@) published 1 November 2001 
Vol. 114(2) published 25 May 2001 Vol. 114(4) published 19 December 2001 


WASHINGTON 
PRINTED FOR THE SOCIETY 


EDITOR 


C. BRIAN ROBBINS 


ASSOCIATE EDITORS 


Classical Languages Invertebrates 

FREDERICK M. BAYER STEPHEN L. GARDINER 
FRANK D. FERRARI 
RAFAEL LEMAITRE 

Plants Vertebrates 

DAvID B. LELLINGER GARY R. GRAVES 
CAROLE C. BALDWIN 

Insects 


WAYNE N. MATHIS 


All correspondence should be addressed to the 
Biological Society of Washington, Smithsonian Institution 
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OFFICERS AND COUNCIL 
of the 
BIOLOGICAL SOCIETY OF WASHINGTON 
FOR 2000-2001 


OFFICERS 
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Elected Members 
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=. 


TABLE OF CONTENTS 


Volume 114 


Anderson, Robert P., and Charles O. Handley, Jr. A new species of three-toed sloth (Mammalia: 
Xenarthra) from Panama, with a review of the genus Bradypus .............0ecce cece eee e eee eeeees 


Assmann, Michael, Rob W. M. van Soest, and Matthias K6ck. Description of Agelas cerebrum, a 
new species and re-description of A. dilatata (Porifera) ............ 6... cece cece cence cee ese eee e eens 


Bannikov, Alexandre F, and James C. Tyler. A new species of the luvarid fish genus ¢Avitoluvarus 
(Acanthuroidei: Perciformes) from the Eocene of the Caucasus in southwest Russia ............ 


Bayer, Frederick M. New species of Calyptrophora (Coelenterata: Octocorallia: Primnoidae) from 
themwecStchmypantonthe AtlantictOCean -s.sssa--eeeee eens aerate se Oeerectee eee een cee see eeeeeraa-ce 


Bayer, Frederick M., and Ian G. Macintyre. The mineral component of the axis and holdfast of 
some gorgonacean octocorals (Coelenterata: Anthozoa), with special reference to the family 
(GOnBOnlIGRS Y sogaaeees ann secs deve pOSne haa aaa eor ida lee rtre scents aa ee arc aan MRS nar strane Leet 


Btazewicz-Paszkowycz, Magdalena, and Richard W. Heard. Observations on Cumacea (Malacos- 
traca: Peracarida) from Antarctic and subantarctic waters. I. Ekleptostylis debroyeri (Diastylidae), 
a new species from waters off the Antarctic Peninsula .................. cece eee eee cece eee ceeeeeeeeee 


Boyko, Christopher B., and Jason D. Williams. A review of Pseudionella Shiino, 1949 (Crustacea: 
Isopoda: Bopyridae), with the description of a new species parasitic on Calcinus hermit crabs 
LT UMMA Ue Taal Say GLY aroetcee epetscee ee yetoree seer a ea ctoee enscoe stars che cps evan ole eroyetanstap ale evcTore oiafa: eel uc wrt ciale onan yaeteialgeotemta tate es 

Cairns, Stephen D. Studies on western Atlantic Octocorallia (Coelenterata: Anthozoa). Part 1: The 
genus Chrysogorgia Duchassaing & Michelotti, 1864 ................ 2. cece cece cece eee e eee eee eeeeeees 

Campos, Martha A. A new genus and species of freshwater crab from Colombia (Crustacea: 
Decapoda wesendothelphusid ae). sk yy sak ctaclots cle lets oeversiers eget e ravines Sia tlds Slejo ate wre abelapoie euslerateverefeveucreyerebatons 

Capa, Maria, Guillermo San Martin, and Eduardo Lopez. Description of a new species of Para- 
Sphacrosyllisi(Rolychaetas Sillidae: Syillamae)) 2.5. cnc. ee eee eee secs eee ees asec ere see seeeeae 

Carleton, Michael D., Steven M. Goodman, and Daniel Rakotondravony. A new species of tufted- 
tailed rat, genus Eleurus (Muridae: Nesomyinae), from western Madagascar, with notes on the 
CSU OWMNIOM Ct Boer ai VT  teoaeo we croseds coeator cote a aaoreaRA eno a comme etoocdecemoeie ora ccrer soo Snes 

Carrera-Parra, Luis E Recognition of Cenogenus Chamberlin, 1919 (Polychaeta: Lumbrineridae) 
DAGEG! Gin {Hiy/OS TESTER eaceasodcenc sadn dod saden Deaoeeees ames odors sear emanasesotuuceanasuensesesuceosean 

Carvalho, Marcelo R. de. A new species of electric ray, Narcine leoparda, from the tropical eastern 
Pacific Ocean (Chondrichthyes: Torpediniformes: Narcinidae) ..................0eeeeeeeeeee eee eeees 

Chang, Cheon Young, and Teruo Ishida. Two new species of the Canthocamptus mirabilis group 
(Copepoda: Harpacticoida: Canthocamptidae) from South Korea .....................2222..220 eee 

Coleman, Charles Oliver, and Judith Kauffeldt. Antarctodius rauscherti, a new species (Crustacea: 
Amphipoda: Ochlesidae) from the Antarctic Ocean .................00 ccc e cece ee cence cence eneeeeees 

Cooper, John E. Cambarus (Puncticambarus) hobbsorum, a new crayfish (Decapoda: Cambaridae) 
iixonm INKoyatal Ceico) bine tM snedeceses ae pe oeae one sem Onda tedas aban neceen cic mae weerea ns se areent mame onias aoe nes 

Corace, R. Gregory, I, Neil Cumberlidge, and Rolf Garms. A new species of freshwater crab 
(Brachyura: Potamoidea: Potamonautidae) from the Ruwenzori region of western Uganda, East 
INTEICEI.. iS ely MOOR INO BROS oO es SUES Tatts Renee Cee reat Oe ee Hee ee Se ST Se EIS OTe 

Curini-Galletti, Marco, Gavino Oggiano, and Marco Casu. New Unguiphora (Platyhelminthes: 
ATO SST ACAD) eT COTATI CLA eres eccrsre cele ater oes Tores oc eae ae ee TE ers assim wis ean TO es tes env Se NG Re in ee 

Cutler, Edward B., Harlan K. Dean, and José I. Saiz-Salinas. Sipuncula from Antarctic waters ... 

Diaz-Castafieda, Victoria, and Guillermo San Martin. Syllidae (Polychaeta) from San Quintin la- 
goon, Baja California, Mexico, with the Gescription of a new genus ...................--.2+--+++-- 

Espinasa, Luis, and Monika Baker Alpheis. A new species of the genus Anelpistina (Insecta: 
Zygentoma: Nicoletiidae) from the Biosphere Reserve Sierra de Huautla ......................... 

Espinosa-Pérez, Ma. del Carmen, and Michel E. Hendrickx. A new species of Exosphaeroma 
Stebbing (Crustacea: Isopoda: Sphaeromatidae) from the Pacific coast of Mexico ............... 

Flint, Oliver S., Jr. Diplectroninae of Sri Lanka (Trichoptera: Hydropsychidae) ..................... 

Fukuoka, Kouki, and Masaaki Murano. Telacanthomysis, a new genus, for Acanthomysis colum- 
biae, and redescription of Columbiaemysis ignota (Crustacea: Mysidacea: Mysidae) ............ 

Galil, Bella S., and Raymond B. Manning. A new geryonid crab from the Amirante Basin, western 
Indian Ocean (Crustacea: Decapoda: Brachyura) .................e eee e cece cece eee cece eee eee e es 

Gardiner, Stephen L., Erin McMullin, and Charles R. Fisher. Seepiophila jonesi, a new genus and 
species of vestimentiferan tube worm (Annelida: Pogonophora) from hydrocarbon seep com- 
MIME S) mnsthe Gullof WIEXa CO; SHEE a the Macc smo tac ben see oats eile a alee obi Ne eee site oth datent apatd «e 


359-366 


579-588 


367-380 


309-345 


907-917 


649-659 


746-787 


938—943 


280-284 


972-987 


720-724 


561-573 


667-679 


427-434 


152-161 


178-187 


737-7145 
861-880 


708-719 


489—496 


640-648 
91-103 


197—206 


104—108 


694-707 


Vi 


Gomez, Samuel. A new species of Onychocamptus Daday, 1903 (Copepoda: Harpacticoida: Lao- 
phontidae) tromimorthwestems Vicxic ON te-eeeeren os Peeeee eee eee sere ease ee Eee eer eee eee a eeeCE eee eeeeee 
Gomez, Samuel, and Sybille Seifried. A new species of Ectinosoma Boeck, 1865 (Copepoda: 
Harpacticoida: Ectinosomatidae) from northwestern Mexico ...............222+-eeeee eee ec eee eee eees 
Graves, Gary R. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 9. Confirmation of the 
hybrid origin of Coeligena purpurea Gould, 1854 .............. 2... e eee cece eee eee eee eee eee eeneeees 
Hershler, Robert, and Daniel L. Gustafson. First record for springsnails (Mollusca: Hydrobiidae: 
Pyrgulopsis) from the northern Rocky Mountains ..................2...csse eee ecceeee eee ee eeeeeeeeees 
Hilbig, Brigitte, and Dieter Fiege. A new species of Dorvilleidae (Annelida: Polychaeta) from a 
coldiseep, site unithe northeast Racihicgs sare a a-a-eeee eee eee reer ee eee ec eee eee eee 
Ho, H.-P, H.-P. Yu, and Peter K. L. Ng. The Indo-Pacific Pilumnidae XIV. On a new species of 
Actumnus (Crustacea: Decapoda: Brachyura) from Taiwan ....................200eeeeee sees teen eee 
Ho, Ju-shey, and Samuel Gomez. Redescription of Lepeophtheirus marginatus Bere, 1936 (Co- 
pepoda: Caligidae) and relegation of L. christianensis Wilson, 1944 and L. orbicularis Shiino, 
MO GOEAS! WES SWIOMIVIIS Tye do. ee os x eee ener ator oreo onto s See nie oe nie ernie SES a Se EIS «ae EOC EER EEEE 
Hooge, Matthew D., and Seth Tyler Interstitial acoels (Platyhelminthes: Acoela) from 
Bermuda fs.).ctee Uh ase Case ae Se A Bad Oe AS 
Ivanenko, V. N., Frank D. Ferrari, and A. V. Smurov. Nauplii and copepodids of Scottomyzon 
gibberum (Copepoda: Siphonostomatoida: Scottomyzontidae, a new family), a symbiont of As- 
térias rubens:(ASteroi1dea) ie eeis. 5 2s. oe eee eas aoe Ee See ee ee eee 
Iwata, Fumio. Nipponnemertes fernaldi, a new species of swimming monostiliferous hoplonemer- 
tean from the San Juan Archipelago, Washington, U.S.A. .............. 2... c cee e eee eee eee eee 


Johnson, G. David, Johnson Seeto, and Richard H. Rosenblatt. Parmops echinatus, a new species 
of flashlight fish (Beryciformes: Anomalopidae) from Fiji ..................0. 2c cee cece eee e cece scenes 
Kato, Hisayoshi, and Akihiro Koizumi. A new fossil geryonid crab (Crustacea: Decapoda: Brach- 
yura) from the late Pliocene to early Pleistocene of Japan .................2.2..eceee cece eeee eee eee es 
Kato, Tetsuya, Fredrik Pleijel, and Shunsuke Mawatari. Eulalia gemina (Phyllodocidae: Poly- 
chaeta), a new species from Shirahama, Japan ............2.-. 62 eee eee ec eee eee e cece tent ee eceeeeaees 
Kensley, Brian, and Niel L. Bruce. Redescription of Dynameniscus carinatus (Richardson, 1900) 
(Cmustacea:slsopoda: ‘Sphacromatidae) i <tosn.s) cease aeocer sete seer eee eec mene dese ce aeee eee eee 
Kensley, Brian, and Cleveland P. Hickman, Jr. A new species of Calaxius Sakai & de Saint Laurent, 
1989, from the Galapagos Islands (Crustacea: Decapoda: Axiidae) .................02--0ee eee ee eee 
Kim, Il-Hoi. Foliomolgus cucullus, a new genus and species of Clausidiidae (Crustacea: Copepoda: 
Poecilostomatoida) associated with a polychaete in Korea ...................02eece eee eee eee eeeeeeeee 
Komatsu, Hironori, and Masatsune Takeda. A new leucosiid crab of the genus Nursia Leach, 1817 
from Vietnam (Crustacea: Decapoda: Brachyura) with redescription of N. mimetica Nobili, 
VQ OG oe) ced Ps BOE ae EEE I I AES GAN IOS SISA 58 SGU AS 
Konishi, Kooichi. First record of larvae of the rare mud shrimp Naushonia Kingsley (Crustacea: 
Decapoda Eaomediudae) tiromeAsiam waters eaenes ete eee see ee ee eee eee eee tee ee ee eee eee eee 
Kornicker, Louis S. Exopod and protopodial endite III of the maxillae of Sarsiellinae (Crustacea: 
Ostracoda: My odocopa)l te I4. see ee Fe as Sh ee ee oe ee eee 
Kornicker, Louis S. Cypridina olimoblonga Kornicker, a new name for the junior primary hom- 
onym Cypridina oblonga Jones & Kirkby, 1874, and clarification of its authorship, and the 
authorship of the Paleozoic genera Cypridinella, Cypridellina, Sulcuna, Rhombina, and Offa 
(Crustacea: Ostracoda). <25.4 -Sesat assessed ccs sate sec Aeoe eae ONE EOS Soe SOE eT SSN De PS ee 
Lambert, Philip, and Kathryn L. Oliver. Pseudothyone levini, a new species of sea cucumber 
(Echinodermata: Holothuroidea) from the northeastern Pacific ...................02.0ecee cece cece 
Lemaitre, Rafael, Julio Garcia-Gomez, Richard v. Sternberg, and Néstor H. Campos. A new genus 
and a new species of crab of the family Goneplacidae MacCleay, 1838 (Crustacea: Decapoda: 
Brachyura) from the tropical western Atlantic ...................cesceeeee seen eect eee e eee e eee eeeeeeeees 
Le6n-Gonzalez, Jesis Angel de, and Vivianne Solis-Weiss. Two new species of Nereis (Polychaeta: 
INereididae) siromethes Mexican’ Paci Canes eeenene a eee eee eee eee eee 
Le6n-Gonzalez, Jesis Angel de, Vivianne Solis-Weiss, and Veronica Valadez-Rocha. Two new 
species of Platynereis (Polychaeta: Nereididae) from eastern Mexican shores ..................-- 
Liao, Yulin, and David L. Pawson. Dendrochirote and dactylochirote sea cucumbers (Echinoder- 
mata: Holothuroidea) of China, with descriptions of eight new species ................-++-++++++++ 
Lindner, Alberto, and Alvaro E. Migotto. Merotrichous isorhiza, a nematocyst new to the Cam- 
panulariidae (Cnidaria: Hydrozoa), and its relevance for the classification of Cnidae ............ 
Lucas, Spencer G., and Robert J. Emry. Sharamynodon (Mammalia: Perissodactyla) from the Eo- 
cene of the Ily basin, Kazakstan and the antiquity of Asian amynodonts .......................--- 


262-274 


207-218 


41-50 


297-308 


396-402 


145-151 


887-906 


414-426 


237-261 


833-857 


497-500 


172-177 


381-388 


188-196 


484—488 


660—666 


599-604 


611-617 


680-685 


686-693 


589-598 


951—963 


881-886 


389-395 


58-90 


825-992 


517-525 


Lucas, Spencer G., Luke T: Holbrook, and Robert J. Emry. The lophialetid ceratomorph Eoletes 
(Mammalia) from the Eocene of the Zaysan Basin, Kazakstan ................:eeecceeeeeee cece eens 


McDermott, John J. Symbionts of the hermit crab Pagurus longicarpus Say, 1817 (Decapoda: 
Anomura): New observations from New Jersey waters and a review of all known 
IRS ATO IN MOS 5 areata dosade pocae dao auane Shoo on ocanRe pace ose He Guero sone ao on tao SROnD Omen San conocer corer 


McLaughlin, Patsy A., and Rafael Lemaitre. Revision of Pylopagurus and Tomopagurus (Crusta- 
cea: Decapoda: Paguridae), with descriptions of new genera and species. Part VI. Pylopagurus 
A. Milne-Edwards & Bouvier, 1891, Haigia McLaughlin, 1981, and Pylopaguridium, a new 
ITS TMULS MSE tae oe on Voinis erates a SyAlsioisin sia laid swchovsusiy Agate cis hcsielsye PRUE RESIS he pera eo eee oles Melee cher bcd cecal 


Mitsuhashi, Masako, Takahiro Fujino, and Masatsune Takeda. A new pontoniine shrimp of the 
genus Corallocaris Stimpson, 1860 (Crustacea: Decapoda: Palaemonidae) from the Ryukyu 
STAN Starr eit cae s ee tice Nucci eR lacie Gia clears be mn sesisis macinag shag s stones unde ct umueee aecae eet 


Morer-Chauviré, Cécile. The systematic position of the genus Basityto Mlikovsky, 1998 (Aves: 
(Giruiiitomm©es:. GiniGki)\. cosvapcusodcds or nba dune amenen ap Roan tina AaNen On ta AE ASS GME area NARS Sa pSe REA art nnaT 


Murano, Masaaki. The genus Pseudoxomysis (Crustacea: Mysidacea: Mysidae: Leptomysini), with 
description of a new species from the Timor Sea ................. cee cc cee een eee e eee eeeeeeceeeeeenes 


Negreiros-Fransozo, Maria Lucia, and Adilson Fransozo. Larval development of Epialtus bituber- 
culatus H. Milne Edwards, 1834 (Crustacea: Decapoda: Brachyura: Majidae) with comments on 
majid larvae from the southwestern Atlantic ............. 0. cc cece cece cece eee e ee een n ete eeeecenneseanes 

Nogueira, J. M. de M., and A. C. Z. Amaral. New terebellids (Polychaeta: Terebellidae) living in 
colonies of a stony coral in the state of Sao Paulo, Brazil ................... 0. ccc cece cece eee e cece ees 

Oliveira, José Carlos de, Richard P. Vari, and Carl J. Ferraris, Jr. A new species of ““Whale Catfish”’ 
(Siluriformes: Cetopsidae) from the western portions of the Amazon basin ....................... 

Olson, Storrs L., and David B. Wingate. A new species of large flightless rail of the Rallus 
longirostris/elegans complex (Aves: Rallidae) from the late Pleistocene of Bermuda ............ 

Opresko, Dennis M. New species of antipatharians (Cnidaria: Anthozoa) from Madeira, with the 
SRPMS VASA OL A) TEN JXSTUG Guocagcocansasarouctancdusananadooneapocsnaueunaasasoonosueadouscsanoccddos 

Opresko, Dennis M., and Rosemarie C. Baron-Szabo. Reevaluation of Tropidopathes saliciformis 
Silberfeld: A hydroid originally identified as an antipatharian coral ...................2.++--eeeeeee 

Orr, James Wilder, and Morgan S. Busby. Prognatholiparis ptychomandibularis, a new genus and 
species of the fish family Liparidae (Teleostei: Scorpaeniformes) from the Aleutian Islands, 
JNIDSEA 25 c0008do0a0 daet ES OHOD TOSI RRSO POCO TOC OE Me eG uO Ss Soe ohne Aneta Steer ene a Seno. ast arse met scree Secs 

Osawa, Masayuki, and Yoshihisa Fujita. A new species of the genus Neopetrolisthes Miyake, 1937 
(Crustacea: Decapoda: Porcellanidae) from the Ryukyu Islands, southwestern Japan ............ 

Parapar, Julio. Revision of five species referred to Myriochele and Galathowenia (Polychaeta: 
Oweniidae) from the Antarctic Seas based upon type material ......................022 222 e seen eee 

Robinson, Harold. A new species of Ageratina from Chiapas, Mexico (Eupatorieae: 
AGISTACEAS)): “sass pdosdasodsso oe BER BEAR e aac ROAD TOTES Cee THs Dee ESI Hues Dea REE On a are ark eer cc Tea 

Robinson, Harold. New species of Fleischmannia from Panama and Andean South America (As- 
TSTACCAS MEM PALOTICAES))| jiercyacers erase cree vials seeieiers orsiGie ates einie ete Miedo wsis cleame ain cinm alee aulajevnia ule srstere evelbia sie Selo sle ares 

Rodriguez, Gilberto. New species and records of pseudothelphusid crabs (Crustacea: Brachyura) 
from Central America in the Museum of Natural History of Tulane University .................. 

Rom4an-Contreras, Ramiro, and Roland Bourdon. Probopyrus insularis, a new species (Isopoda: 
Bopyridae), a parasite of Macrobrachium faustinum (de Sassure, 1857) (Decapoda: Palaemoni- 
dae), with criteria to differentiate species of Probopyrus ........-..12eccc eect nee eee eee teen eens 

Sara, Michele, Patricia G6mez, and Antonio Sara. East Pacific Mexican Tethya (Porifera: Demo- 
spongiae) with descriptions of five Mew SPeCieS ...........-.. eee ee eee eee eee cette e eee e eee e eee ees 

Scelzo, Marcelo A. First record of the portonid crab Arenaeus cribrarius (Lemark, 1818) (Crus- 
tacea: Brachyura: Portunidae) in marine waters of Argentina .....................2......eee eee eee 

Seo, In Soon, and Kyung Sook Lee. Haplostoma kimi, a new species and a redescription of 
Haplostomella halocynthiae (Fukui) from Korea (Copepoda: Cyclopoida: Ascidicolidae) ....... 

Shimomura, Michitaka, and Shunsuke E Mawatari. A new asellote isopod of the genus Santia 
Sivertsen & Holthuis, 1980 (Crustacea: Isopoda: Asellota: Santiidae) from Japan ............... 

Sgrensen, Martin V. On the rotifer fauna of Bermuda, including notes on the associated meiofauna 
and the description of a new species of Encentrum (Rotifera: Ploima: Dicranophoridae) ....... 

Suarez, William, and Storrs L. Olson. A remarkable new species of small falcon from the Quater- 
NanytOm CubacAvess Halcomidac: HalGO)). wake cneceoaeeeientars oni agen «ae telsie eee haieisoate ao ee oa eee sis 


Suarez, William, and Storrs L. Olson. Further characterization of Caracara creightoni Brodkorb 
based on fossils from the Quaternary of Cuba (Aves: Falconidae) ................-.-.22+-2eeeee ee 


Vil 


988-992 


624-639 


444-483 


944—950 


964-971 


887-896 


120-138 


285-296 


574-578 


509-516 


349-358 


788-793 


51-57 


162-171 


403-413 


526-528 


529-556 


435-443 


918-928 


794-821 


605-610 


229-236 


929-937 


725-736 


34-40 


501—508 


Vill 


Suarez-Morales, Edwardo, and Cristina Dias. A new species of Monstrilla (Crustacea: Copepoda: 
Monstrilloida) from Brazil with notes on M. brevicornis Isaac. ................000e cece cence cece eee 


Waller, Thomas R. Dhondtichlamys, a new name for Microchlamys Sobetski, 1977 (Mollusca: Biv- 
alvia: Pectinidae), preoccupied by Microchlamys Cockerell, 1911 (Rhizopoda: Arcellinida) ...... 


Werding, Bernd. Description of two new species of Polyonyx Stimpson, 1858 from the Indo-West 
Pacific, with a key to the species of the Polyonyx sinensis group (Crustacea: Decapoda: 
Porcellanidae) 


Wicksten, Mary K., and Rita Vargas. A new species of Thor Kingsley, 1878 (Crustacea: Decapoda: 
Caridea: Hippolytidae) from the tropical eastern Pacific ................ ss see e eee eee eee eee t eee e eens 


Xie, Zhicai, Yanling Liang, and Ji Wang. Fridericia nanningensis, a new terrestrial enchytraeid 
species (Oligochaeta) from southwestern China ................... cee eee eee cece cect eee e eee eeeeeeeees 


Yeo, Darren C. J. A new cavernicolous species of freshwater crab (Crustacea: Brachyura: Potam- 
idae) from Palau Tioman, peninsular Malaysia 


219-228 


858-860 


109-119 


139-144 


275-279 


618-623 


INDEX TO NEW TAXA 


Volume 114 


(New taxa are indicated in italics; new combinations designated n.c.) 
PORIFERA 


PNGSMENS COT DI PU aa ta Ne re ee 


Tre ny CAR 70.5 1S fe ee a ee Eh a Te rn 
FEL CONG G CLIT Clase aa ca ERT PI LON Tu st SO ee Sa I ne PD os FE A aed CO 
CNEL Ta ae 2 RN an in OD rs in Ee 
|OIRORGE LIT hy a ee Pe ee eee eee eee 
OCHA axe SE ORS ee, STE SE coy REM 9 Cee gs NOW Pee Oe NA Ea AU ICS 


(Coa DAUR ODL NO NY CHAT LLG SN OE A a Pe OE eae 
SGU: Se ee eee eee ee ee eee eS 
[ELIS GHED  O  enepe eeee 


CGhimySOPOnsIaAMReKGend Opin (ee ee ee ee 


TE IAN EXCEL ANG. Et a a 
(GOL TTLUG Ol Cee a ere tea ean at oe I ee pe ease 
OUTAGES Se ta ele ae eR eNO C1 a 9 SEE ee ST SRE eee UN eae teat at 


ANIONS BAY OY AC 5 ee Se Oe Sr Ee eee eee ee 
OLA CLC: Np ee Re RR ee nC ree ee 


ANGERROSOZEL GR, ELIE a ee eee 
COUPLING ee en ge re a a en i ee 


Lap) LO pas Chana aye CLUE Cae eta a see NON GE Ee a ee 


IN eames ae 
TCL CNN Ne RAN MNS al isch De a out ra Me oa SL I Le a oh abe pes He Bean ey 


LPP EY ONG) ORNS H SVE AAW ELIT (0): 5 eee ree Se ee 


IESTi@) [OT SHEA GT TLE CL CTS US pm one sere cE Ae a a ae 
IN POP OMDINS ITN Tet Say C77 Cl ee ee era ane ewe eerie ee eee Sere ee ee 


FEST GG GTN EGURTNM CLS TG CL Ca Cag a eer eta a Si aA Ba cl Recon ae ele ee he SS 


ANNELIDA 
Polychaeta 


GLC C'S CN eR OMNI a Sioa re a aa EA Be AS AL a re OE a cr mc ON Dl 


ESM AU AS27007 eg eae a ne I IL a acces 02s I BOR PRES USES 


Grubeosyllisymediodentata msc.) es ie Se ee a 


MOVSQNG: 2. Auf Se a ee ee EEE ————————eeeee 286 


DiSCLIOSG): Bevas 5 eat ete he Se ee ee eee 288 
INGLES \COSOGCY eee ee cos Ne ee ee a eee ee 881 
DA UCI GI RA SE AE 2 TS a Ee 882 
Parasphaerosyilis (nQlinia lita. ee 280 
Platynereisvhuichinesac’ (= 2a eee 389 
TULCTONGLG! ee Ea a NE ee ARS De Pe 391 
RarougiaiOreSOnensis | qs _ eS ee ee ee 398 
Stieblosomaalisobranchiatum” 22 = eee 292 
Oligochaeta 
Rndenclalvannincensis = ee 276 
Pogonophora 
SCO PDLOP PAUL aS oP a al ty see Se a 697 
OTL CS Dn saan Ba Raa eNO R ea Pa NR a er op 697 
ARTHROPODA 
Crustacea 
AGWIMINUS (QIWANCHSIS® = 80 ee ee eee 145 
Antarclogiusi7 aus Chertin. <2 ee eee 427 
ACRGGUGS 22008 eek AS, SS ie Bienes fan Sl eal eT 938 
COS GHGTCTUSES, <a Be lle 939 
Calaxiusealapagensis= === ee ee ee 484 
Cambanis (Puncticambarus) hobbsorumt a 153 
Canthocamptusineunviseiosus. 222). 2 et ee eee 672 
DADC CIS US je ee eke Ba Ia Ss a Bi aD Be a en 668 
Ghaceonuconernt jt ee ee ee 104 
MGLSUSH ING te 2 ee 173 
Coralliocanisuinidens) “22822 aie Sa ee ee ee 944 
(Ghyyjoraaliinen, @liswovolo\oravercy, TVe\w7 TO Na NYS) 686 
ECtimosoma. mexicanum) = 2s.) ee ee eee 207 
Ekleptostylisidebroyert> 2.3 i nn 908 
INOMAtQlIN CS on -8 =: SN ee ne ee ee ee ee 914 
Exosphaeromabyuscat. 2222 ee ee 641 
PF OVWOIMIOUV GUIS © 2 tone oe wal Ree ne 2 ee ee 660 
CUCU AS Se EN hd SN ee 661 
BVeapo) Stub ea ACEP PGT aa el ae 229 
WOMGr al Ora ° jose kc ti Ba Rodan EE SB clit aes ue eaten ee 618 
Monstrella\ pustuilatas nc 0 a 219 
INGHES PUG nes 8 APS Ls, ER A ee 952 
VOSSU yc a a ee i 954 
Neopetrolisthtes spinatus. <tr 163 
INUUSIA AOUUILOLGE. ga a ee 599 
Onychocamptus (ramisaustralis, =. 2 ee ee 264 


|G. YA) 612 Gay (121 | eee ee ee cc et Or SS I 0 te oe 112 


EEA UCT US pe te uo eR en oe ane seen YE 
RO Latin OMAUTS Serst6 kW CUTU te eee eee re ee er eo een ee ee ee 
ROTA AA GATCITNUISHVLEL COULCTU STIS | tea I a re ee Ore ee 
TPYROUBYOYDY RUNS OSU CIAL a a eee 


TPrsysxuna iyo vel lei Tam eee sca eg ahaa Ne et ce ee cae rm Sc eR OU <a eR Otel he ea 
CATCH IS Fk ea IS rs BD a Re eee Ra RRP een, 5 ROUEN ett cee ere ete 


SUL GH Gy ADT, SUG GUE 72 GUS Co eager arma oN ak SAI Tr SO Ear a Sc ne tS RO 
LP ASS ACG we ASS « LAH IAS i a RR a ee ee 


SEAS) COED CUSOL gL CULT gum om tte Dee Ee BSP Lz BN a SOR Be Da A a dt a eel a Slee 
TL ELCUIGRSLCLT TL Ugg Pee eee Oe eee SIRE ace tal Se rn ee eee Ne AU ee 


LPAI KOY BEBY EAU UNS SONG ee tN Le SN a La a ee ee ee meen ee ee Oe ae 
UPL ESO DIOR ss a I I ee 


PS AI ULEAM LO UTS CL OS Cla te ete sem ep I A A 3 Ta Ra i ene ee nae eee 
S\COi ROMY AOU ONG Ys 5s ee Oe ee eee 


TE QUOTA TOTS SIS at a Ne Ae Oe ee Ree ed a 
SCO LM Ten NTA at Gp teed il oa ets ll bg a ice gs IS aN ates en cal Ny heath aloes) oa PA 


WIRING Ta OG OT LCT US US fase ae tae aed a ee EN a aa ee Be ree ee 


PANNE Ne I PS fe1NN ACL OYC1 CL OU se a Bt NT a ne 


ISH USANA, OSS ec Ae 
LOVES US ty Lea eee an Ok EN ES PS SEE Ss SEEN Oe ES AE Se LE 1 eee re 
IEICE: yy Tyme esa Je Se I a See ee er 
(UPI OLDE NAV SAS 1 Cs eS Ss ARE 5 A ee A ee 


PNCIMITO CUCM SC HEIL CTS US) ian tet rw ce ti tae sr ee Ns A ace na eae een 
Tsophyilophorusmewssubs emu sys Weeks ee eee 


ING Clan OTM GAMUT PUTT UL TT pa ae a et a ee re 


MU OH ERPDU PUT COPUTLGL AIAN tere mt ender SUE ee, va eee eS ee eee 
ESULLOUE S TS act orc emirates ON UE eral Ni rosea AE Se eo UE els Sn ee eal OO anne, ieee 


CHORDATA 


Pisces 


PANIC O MUN AGIISHOOC CII CLUS eee eee ee ere aN Os ea oe a Oe a re ee 
(CSCO SUS WP) 2111, CL meee ea tes UN NT 0) Ney wn wg Rae SOL ed eee Na ees ee ee en 


INA CUT O TCLT;L Cr aetna se ec Rn ee Ie rt 


Xi 


Xi 


Pari OPS CCH UIA US = re ee en 497 
Te on AN Op eA S a ee a ce 51 
pliychomandibulant’ = ee eee 52 
Aves 
Baleanicaymummrmela, ms Cys oe ee te 964 
CO Fig 7O CPG Da Na he cs a ee ele tere 35 
RANISHRECES SES PM te oe ee ae ete ie ple han ene te tle ep Syl il 
Mammalia 
Brady PUS Py QTae Us, ee ee a ee Se 17 
VU UUEUUS a1 BS 170 yy A I a a ea pel rn 974 
PLANTAE 
AWeerannai(Neosncenclla)inesinijend = 526 
Fleishmannia alihuanucana se ————————E———E————ee— 544 
CL] QUILL CYUS ES a 545 
COTLONUUE oa eae ee et ed 535 
GAViGS Tpit hatt” eaee  e e  e 548 
LOGS OFA Ui a a a ee 2 Pe ere cer eee ot ee ek eerie 535 
hammellit) 2225 ee ee ee 530 
SED LE Wig ae he i, 5 SES Bee en ee 531 
IGLID OU ees sh eee RS ee ee ee ee ee 538 
WL CTOSCCTI OUD OS a a eet ee ee 540 
TLCUVEVLO CTUSUS) ec ic i aha a ne ee 533 
PCT OL AUN na a di a SO Stn a Lvl lect asta 548 
LUT OU peas tS Be dG aa ao oh sie Be 550 
SCL SUC UAUD «cee eM ele 2 a ee 552 
CLIC CES UU creer rene Oe eV ee ny SRO Se SRO ete NOB a te 556 


CANT Ui ee OO Ro Se aR Rea al SEE EOD eee A eS ec 541 


ee 


INFORMATION FOR CONTRIBUTORS 


Content.—The Proceedings of the Biological Society of Washington contains papers bearing 
on systematics in the biological sciences (botany, zoology, and paleontology), and notices of 
business transacted at meetings of the Society. Except at the direction of the Council, only 
manuscripts by Society members will be accepted. Papers are published in English (except for 
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Presentation.—Clarity of presentation, and requirements of taxonomic and nomenclatural 
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“GUIDELINES FOR MANUSCRIPTS for Publications of the BIOLOGICAL SOCIETY OF 
WASHINGTON,” a supplement to Volume 114, number 4, December 2001. Authors are encour- 
aged to consult this article before manuscript preparation. Copies of the article are available 
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inch per column. One ms. page = approximately 0.4 printed page. 


Front cover—from this issue, p. 945. 


CONTENTS 


Merotrichous isorhiza, a nematocyst new to the Campanulariidae (Cnidaria: Hydrozoa), and its rele- 
vance for the classification of Cnidae Alberto Lindner and Alvaro E. Migotto 
Nipponnemertes fernaldi, a new species of swimming monostiliferous hoplonemertean from the San 
Juan Archipelago, Washington, U.S.A. Fumio Iwata 
Dhondtichlamys, a new name for Microchlamys Sobetski, 1977 (Mollusca: Bivalvia: Pectinidae), pre- 
occupied by Microchlamys Cockerell, 1911 (Rhizopoda: Arcellinida) Thomas R. Waller 
Sipuncula from Antarctic waters Edward B. Cutler, Harlan K. Dean, and José I. Saiz-Salinas 
Two new species of Nereis (Polychaeta; Nereididae) from the Mexican Pacific 
Jesus Angel de Le6n-Gonzdlez and Vivianne Solis-Weiss 
The genus Pseudoxomysis (Crustacea: Mysidacea: Mysidae: Leptomysini), with description of a new 
species from the Timor Sea Masaaki Murano 
Redescription of Lepeophtheirus marginatus Bere, 1936 (Copepoda: Caligidae) and relegation of L. 
christianensis Wilson, 1944 and L. orbicularis Shiino, 1960 as its synonyms 
Ju-shey Ho and Samuel Gomez 
Observations on Cumacea (Malacostraca: Peracarida) from Antarctic and subantarctic waters. I. 
Ekleptostylis debroyeri (Diastylidae), a new species from waters off the Antarctic Peninsula 
Magdalena Blazewicz-Paszkowycz and Richard W. Heard 
Probopyrus insularis, a new species (Isopoda: Bopyridae), a parasite of Macrobrachium faustinum (de 
Sassure, 1857) (Decapoda: Palaemonidae), with criteria to differentiate species of Probopyrus 
Ramiro Roman-Contreras and Roland Bourdon 
A new asellote isopod of the genus Santia Sivertsen & Holthuis, 1980 (Crustacea: Isopoda: Asellota: 
Santiidae) from Japan Michitaka Shimomura and Shunsuke F. Mawatari 
A new genus and species of freshwater crab from Colombia (Crustacea: Decapoda: Pseudothelphusi- 
dae) Martha R. Campos 
A new pontoniine shrimp of the genus Corallocaris Stimpson, 1860 (Crustacea: Decapoda: Palaeonidae) 
from the Ryukyu Islands Masako Mitsuhashi, Takahiro Fujino, and Masatsune Takeda 
A new genus and a new species of crab of the family Goneplacidae MacCleay, 1838 (Crustacea: 
Decapoda: Brachyura) from the tropical western Atlantic 
Rafael Lemaitre, Julio Garcia-Go6mez, Richard v. Sternberg, and Néstor H. Campos 
The systematic position of the genus Basityto Mlikovsky, 1998 (Aves: Gruiformes: Gruidae) 
Cécile Morer-Chauviré 
A new species of tufted-tailed rat, genus Eleurus (Muridae: Nesomyinae), from western Madagascar, 
with notes on the distribution of E. myoxinus 
Michael D. Carleton, Steven M. Goodman, and Daniel Rakotondravony 
The lophialetid ceratomorph Eoletes (Mammalia) from the Eocene of the Zaysan Basin, Kazakstan 
Spencer G. Lucas, Luke T. Holbrook, and Robert J. Emry 
Contents, Volume 114 
Index to New Taxa, Volume 114 


825 


833 


858 
861 


881 


887 


897 


907 


918 


929 


938 


944 


951 


964 


QH 
i 
B4x 


Nt 


PROCEEDINGS 


OF THE 


BIOLOGICAL SOCIETY 


OF 
WASHINGTON 


om 
cr 95 we » ) 


LIBRARIES 


GUIDELINES FOR MANUSCRIPTS 
- For Publications of the 


BIOLOGICAL SOCIETY OF WASHINGTON 


Supplement to Volume 114, No. 4 


December 2001 


Proceedings of the Biological Society of Washington 


VOLUME 114 NUMBER 4 
DECEMBER 2001 


send proofs to: 

—Dr. C. Brian Robbins 
—11017 Via Merida 

—lLa Mesa, CA 91941-7381 


Guidelines for manuscripts for publications of the 
Biological Society of Washington 


C. Brian Robbins and David B. Lellinger 


(CBR) 11017 Via Merida, La Mesa, California 91941-7381, U.S.A., 
e-mail: robbins.brian@home.com; 
(DBL) Department of Systematic Biology—Botany, National Museum of 
Natural History, Washington, D.C. 20560-0166 U.S.A., e-mail, 
lellinger.db@nmnh.si.edu 


Abstract.—The abstract should be =3% of the length of the 
text of the article. It should be a concise statement of findings, 
rather than a listing of subjects covered, and should be written as 
a single paragraph and double spaced. Except under unusual cir- 
cumstances, references should not be cited in the abstract. All 
newly described taxa must be given by name in the abstract. 


This paper provides authors with guidelines and examples to aid in 
_ preparing manuscripts for the Biological Society of Washington. The 
required format for manuscripts for the Proceedings and Bulletin is 
presented and followed herein as a model of format and style. However, 
except for the title, author(s), author(s) address(es), abstract, and figure and 
table captions, the final printed papers appear in double-column format 
rather than occupying a complete page as does this manuscript. It is neither 
possible nor desirable to provide detailed examples of every format or style 
item, but we have attempted to address most of the potential questions 
facing an author preparing a manuscript. 

As a general guide, we recommend the sixth edition of Scientific style 
and format: The CBE manual for authors, editors, and publishers (Style 
Manual Committee 1994), available from the Council of Biological Editors, 
Inc., New York. The latest edition is a more comprehensive guide for 
scientific writers than were previous editions. A shorter but useful guide is 


Day (1994) How to write & publish a scientific paper, fourth edition. 


1 


Nw 


Methods 


Manuscript preparation.---Type all manuscripts on letter size (215- by 
280-mm, 8'2- by 11-in.) paper. All parts of the manuscript should be double 
spaced, including text, literature cited, tables, table headings, and figure 
legends, so that reviewers, editors, printers, and proofreaders can make 
essential notations. Use the same font size throughout the manuscript. Do 
not justify the right margin. Allow =3-cm (1% in.) margins on all sides. 

Do not use a title page. Type name, address, and telephone number, and 
e-mail address of the author to whom all correspondence and proof is to 
be sent at the upper left of the first page. (Note to foreign authors: this is 
the place to indicate the name and address of the person in the U.S.A. who 
will read proof and order reprints.) Place page numbers at the upper right 
of all pages except the first and do not use a header. 

Titles should be brief and include key words that will be useful for 
indexing and information retrieval. Include phylum, class, order, and family 
names (separated by a colon and a space [e.g., Crustacea: Copepoda: 
Caligidae]) for species likely to be unfamiliar to many readers. In 
subsequent use of scientific names, abbreviate generic names where 
possible (e.g., “Studies on Botrychium meridionale and B. virginianum’’ ); 
do not abbreviate scientific names in titles or subheadings if they have not 
been previously spelled out in the same title or subheading. Center the title 
and type it in uppercase letters, including underlined scientific names. 

Three spaces below the title, type the name(s) of the author(s) centered 
on the page. Triple-space after the author(s) name(s) and enter their 
addresses. If the first author is not the correspondent, indicate after the 
appropriate authors name with an asterisk (*). At the bottom of page | will 
be an asterisk indicating that this is to be the corresponding author with 
their e-mail address. The format for indicating the address of each author 
is: (JD) John Doe, etc. Then begin the abstract, using a normal paragraph 
indentation as shown on page | (secondary heading—see below) and write 
the entire abstract as a single paragraph. Also, indent five spaces at the 
beginning of each paragraph throughout the text. Double-space after the 
abstract and insert a 5-cm horizontal line. Double space after the line (as 
on page 1) and begin the introductory material. Do not use “‘Introduction” 
as a heading. 

In general, use no more than three levels of organization: primary 
headings (centered); secondary headings (indented from left-hand margin, 
underlined, followed by a period and three dashes); and tertiary headings 
(indented from left-hand margin, ending with a colon). New taxon names 
and associated citation of their taxonomic placement (phylum, class, order, 
family, etc.), and figure and table references are primary headings. (Note: 


» ji. 


3 


unless the higher taxonomic categories are the primary topic, author and 
date citations are not necessary.) 


Systematic Paleontology 
Class Mammalia Linnaeus, 1758 
Order Chiroptera Blumenbach, 1779 
Family Desmodontidae Gill, 1884 
Genus Desmodus Maximilian, 1824 


Desmodus archaeodaptes, new species 
Fig. 1 


Long lists or complex material that is related only indirectly to the 
primary subject should be placed in an appendix at the end of the 
manuscript. Items that are appropriate for appendices include: lists of 
specimens examined; lists of morphometric characters, other morphological 
features, or ecologic characters; derivations of complex mathematical or 
Statistical formulas; and algorithms for computer programs. Appendices 
may be arranged as paragraphs, in tables, or in other formats as appropriate. 
Short lists, single formulas, and material related more directly to the subject 
should be placed in the text. Material not essential to the article should be 
omitted. 

Never break words at the right-hand margin anywhere in the manuscript 
for any reason. This includes hyphenated words and words divided into 
syllables. 

Spell out the entire scientific name the first time a plant or animal is 
mentioned in the text; thereafter, abbreviate generic names (first letter and 
period), except at the beginning of a paragraph or sentence. Include 
- author(s) and date where such is essential to identification of a taxon (such 
aS in a synonymy). If subgeneric names are used, they should be placed 
between the generic and specific names, and be enclosed in parentheses; 
abbreviate them after their initial use, using the first letter of their name. 
Use subspecies or infraspecific names only when they are essential to the 
understanding of the article. Descriptions of new taxa in zoology should 
be spelled out, as: new genus, new species, new subspecies, and new 
combination, or for botanical papers use: gen. nov., sp. nov., subsp. nov., 
or comb. nov. Each of these should be preceded by a comma. Descriptions 
must be in English, except for Latin descriptions or diagnoses in botanical 
papers (fossils excepted). 

Provide the scientific name (generic and specific names) of all organisms 
at first mention of the vernacular name both in abstract and text (even if 
the name appears in the title); use the scientific name of the appropriate 
taxon if a vernacular name is inclusive (e.g., crickets [Gryllidae]). 


Tables and figures.---Because of the cost of publication, tables should 
be used only when large quantities of data must be summarized or trends 
in data illustrated to support inferences. Pertinent data in most small tables 
(two-row or two-column tables) usually can be presented in the text in less 
space and without loss of intelligibility. Consider carefully the presentation 
of non-tabular data in the two-column format of the Proceedings. In many 
instances, a table would be easier to read and type-set. 

Figures and tables should be numbered in the sequence in which they 
are referred to in the text. Reference in text should be in the style: (Fig. 1) 
and (Table 1). Reference to several tables or figures in text should be in 
the style: (Tables 1, 2) and (Figs. 1, 2). Parts of composite figures should 
be labeled with uppercase or lowercase letters, or numbers, and should be 
used to refer to parts of figures both in legends and in text (e.g., Fig. la, 
b, d; Figs. 1-3, 5; or Figs. 1B-D, 2D-F). Then note the desired location 
for each figure and table in pencil in the left margin of the manuscript. 

Because many readers prefer to scan data before reading text, each table 
or figure with its title or legend should be understandable without reference 
to the text. Be sure to include species, localities, and dates of study, if 
appropriate, in addition to a description of the content. Abbreviate words 
in column heads or data identifiers (stub columns) only when necessary. 
Spell out names of months except where space does not permit. When 
abbreviated, and in the Materials section, use three-letter abbreviations 
without periods (e.g., Jan, Jun, Oct). Reference in the text to tabular and 
graphic presentations of data should be in support of inferences, not simply 
to direct the reader to specific tables or graphs. For example, use sentences 
such as “‘Litter size did not seem related to month of onset or duration of 
the reproductive season (Table 1), or “‘Morphometric data for type 
Specimens are given in Table 1,’ but avoid sentences such as “Data on 
litter size, and month of onset and duration of the reproductive season in 
some Spermophilus are presented in Table 1.’’ Do not describe the content 
of tables and figures in text; the need for such description indicates that 
the tables and figures are not understandable by themselves. Data should 
be presented either in graphic (figures) or tabular form, whichever is more 
understandable or economical of space. Figures should be constructed to 
be fully legible and not wasteful of space when reduced to column width 
or page width. Orient multiple graphics having two or three illustrations 
one above the other if reduction to a single column is desired. Illustrations 
that are intended for a full-page presentation should be mounted 
accordingly. A figure that is intended to occupy two successive or facing 
pages must be avoided. 

Type each table on a separate sheet(s), give it a complete, intelligible 
title, and refer to it by number in the text. In titles of tables, italicized and 


5 


Roman type is presented as in the text (i.e., underline taxon names). 
Capitalize only the first word of column heads and items in the secondary 
or data identifier column heads; do not use all uppercase letters in headings. 
Do not use a dash or minus sign in a table to indicate lack of observations 
or tests if the table contains plus and minus signs; otherwise its use is clear 
(= missing data). Footnotes in tables should be kept to a minimum. 
Lowercase letters should be used to denote footnotes except those limited 
to probability; one, two, or three asterisks (*) within the table should be 
used for P < 0.05, 0.01, and 0.001, respectively. 

Provide a legend for each figure and refer to the figures in the text. Type 
all figure legends, double spaced, on a separate sheet at the end of the 
manuscript; the typescript format for figure legends is illustrated in 
Appendix I. In legends, underline only those words or terms (i.e., genus 
and species names) that will be set in italics (as listed elsewhere in this 
guide). 

Place any necessary identifications (e.g., symbols, cover types, scale 
bars, regression formulas) directly on the figure rather than in the figure 
legend. Scale bars should always be included in the figure, not in the figure 
legend. Do not submit figures larger than 215 by 280 mm (8% by 11 in.). 
If original figures are larger than this please send a photographically 
reduced figure. Figures exceeding this size are difficult to mail, handle, and 
store. Prepare line drawings with lines of sufficient width and letters of 
sufficient size to remain legible when the figures are reduced to final page 
or column width (Style Manual Committee 1994). Line art should not be 
prepared for reduction greater that 50%. Graphics produced by coarse dot- 
matrix printers are not satisfactory for reproduction. Legibly mark all 
photographs and figures on the back with the author(s) name(s), figure 
- number, and indicate ““TOP.”’ Use soft pencils (blue preferred) on the backs 
of photographs and drawings, because markings, pencil, and pen 
indentations may show through. Leave a margin at the top of each figure 
Gncluding photographs) of at least 3.85 cm (1% inches). Provide 
photocopies of figures to be sent with the manuscript to reviewers. Rub- 
on or transfer letters are preferred. Submission of photographs (either those 
produced by standard processes or by diffusion-transfer imaging [PMT)]) 
of graphics is recommended. Submitted photographs must not be produced 
by making a photograph through a halftone screen (pre-screened). Mount 
original graphics or photographs on cardstock or other heavy material; use 
the same material when several small graphics are submitted for a 
composite figure. In the case of composite graphics, they must be mounted 
as the author desires. If more than one image is mounted on a figure, it is 
best to cut square edges and mount the images with edges touching but not 
overlapping (note: our printer will add narrow white rules between the 


6 


individual images). Also, try to place the identifying letters or numbers 
inside each individual photograph—at least ¥, in from all edges. The editors 
will not mount photographs. Indicate all cropping instructions clearly on 
the original art (or on the photocopy if there is no room on the original). 
For clarity of presentation, the Biological Society of Washington prefers to 
print photographs in their original submitted size, or a size reduction to fit 
a journal page; consideration of size should be made before preparing a 
composite figure. 

Color figures: We accept reflective copy (color prints) and transparencies, 
although it is usually easier to work with reflective copy. If you are creating 
color figures electronically, you must set your graphics file and output 
device to CMYK mode, not RGB. Color graphic file resolution should be 
300 dpi. An original color print is preferable to a scanned electronic copy. 
Figures that will be reproduced in color should not be mounted on stiff 
board as they will be scanned on a drum scanner and need to remain 
flexible. For multiple color images on the same figure, mount them as close 
together as possible. Our press will add the white rules between images. 

Electronic artwork: We accept line art, halftones, and color figures on 
disk. You must attach a printout of the file list that includes the file name(s), 
size, and kind of file in the graphic folder or directory. Indicate the input 
resolution on any images that have been scanned. We require grayscale 
images to be scanned at 450 dpi. All color figures should be scanned at 
300 dpi. All line art should be scanned at 1200 dpi. Please note that nearly 
all images that are downloaded from the Internet, or that are in JPEG or 
GIF format, will be 72 dpi and not acceptable for the printing process. We 
will also not accept graphics embedded within a text file. Indicate the file 
format of the graphics. We accept TIFF or EPS format. Include screen and 
printer font files for any text that has been added to the figure. PC or 
Macintosh versions of Adobe Postscript fonts should be used. Do not use 
True-Type formats. Do not use system “bitmap” fonts. If you have 
compressed or flattened your files, indicate what compression format was 
used. We can accept graphics on the following disks: 3%" disks, Zip disks, 
Jaz disks, or CD-ROM. 


Results and Discussion 
Style and Usage 


Punctuation.---Use quotation marks sparingly for emphasis or special 
use of a word or term, thus preserving them mainly for actual quotations. 
Use the slant line or solidus (/) only to indicate “‘divided by” or in dental, 
vertebral, or chaetotaxy formulas. 

Underline generic and specific names in the text (underlined words will 


7 


become italic in print). Do not use italic or bold type. Underlined words 
(italics) are discouraged for use other than scientific names and headings. 
However, all terms and symbols in mathematical equations and those used 
to denote statistical tests should be underlined. Also, some words that need 
to be emphasized can be underlined and should be so indicated (underlined, 
not in italics). 

Hyphenate compounds used as adjectives (e.g., 3-year-old male, 77-day 
gestation period, 10- by 10-trap grid, home-range area, life-history strategy, 
0.5-m plot). The same holds for adverbs (e.g., well-developed dentition) 
except those ending in “‘ly.”” However, to emphasize the true subject, use 
prepositions to avoid strings of modifiers (e.g., estimates of home-range 
size, not home-range-size estimates). Hyphenation in formation of 
compounds is complex (e.g., ““semi-independent’’ but “‘semiarboreal”’ and 
“mid-July” but “‘midday’’); use of Webster’s Third New International 
Dictionary, Unabridged is advised. 

Always use serial commas, including that preceding the conjunction 
(mice, voles, and shrews). Use no punctuation between state and zip code 
in addresses (address line of these guidelines is an example). Do not use 
‘“‘back-to-back”’ parentheses () (), as for citing a reference and referring 
to a table in text; use (Smith 1984, Table 1) not (Smith 1984) (Table 1). 
To separate a parenthetical statement from a reference in the text, use a 
l-em dash (three dashes on typewriter) if the authority is cited in support 
of statement as in (... as commonly seen in Peromyscus---Smith 1984); 
otherwise, separate with a semicolon as in (.. as commonly seen in deer 
mice [Peromyscus leucopus; Wolff 1988]). 


Abbreviations, spelling, and miscellaneous items.---Paragraphs and 
sentences should not begin with an abbreviation (such as P. maniculatus 


was taken ...). Do not use contrived acronyms or mnemonics for names 
of localities, study areas, morphological characteristics, governmental 
agencies, physiological parameters, statistical tests, or most other items. 
Acronyms for museums, standard abbreviations for protein and enzyme 
loci, and symbols used in mathematical equations are acceptable if 
referenced or defined at first use (words should be used for the latter when 
not used in a mathematical sense; e.g., ““The area is mr’, but the radius is 
shorter than the diameter’’). 

Spelling and use of words should be in accordance with Webster’s Third 
New International Dictionary, Unabridged. Use of words not included 
therein should be avoided, but newly coined words and foreign words may 
be used sparingly if precisely defined at first use. Scientific terms should 
follow recent specialized dictionaries and glossaries. Refrain from using 
words in other than their standard meanings. 


Use only “‘male’’ and “female” to distinguish the two sexes. Avoid 
terms such as rams, ewes, bucks, or other similar names. 

Numbers and mathematics.---Use decimals rather than fractions except 
in equations. Decimals not preceded by a whole number should always be 
preceded by a zero (0.75) in text, tables, and figures. 

In the text (antroduction, discussion, conclusions, etc.) and in titles, use 
numerals for numbers greater than nine except when starting a sentence 
(associated units should then not be abbreviated: ““Eleven minutes,’’ but 
**About 11 min’’; “‘Fifteen percent,” but ““More than 15%’’). For numbers 
one through nine, use words except when used with units of measure (6 
mm), time (3 days, 3 summers, 4 years), but not enumeration (five dugongs, 
two crabs, seven observations). Also, use numerals for all items in a series 
that includes at least one number greater than nine (1 dik-dik, 7 numbats, 
and 19 slow lorises). Treat ordinal numbers in the same manner as cardinal 
numbers (first panda, 14th deer mouse, Ist month, 6th min, fourth trial). 

In the specimens examined and description or diagnosis sections, use 
numerals throughout. 

Avoid the use of numbered sentences or phrases in text. Reference to 
specific items by number in text is acceptable but names of items should 
be lowercase (e.g., day 1, experiment 4, setiger 5, grid 6, site 1, coxa 3, 
segment 7). Do not refer to individual animals either by name or field- 
catalog number; reference to individual animals tends to indicate that the 
material being presented is anecdotal. 

Use commas in numbers of five digits or more (e.g., 10,000 and 
100,000). The exceptions are field and catalog numbers of museum 
specimens, and pagination in references. 

Use a colon (not a solidus [/]) to express ratios (e.g., 1:3.2, males: 
females). Do not present the numbers of males and females in a total 
sample (e.g., 15:48) as sex ratios but calculate the true ratio and give the 


sample size. For example, “... 1:3.2 (n = 63) in favor of females.”’ Use 
< and > for “less than”’ and “‘greater than”? with numerals (“<5 g,” not 
“less than 5 g’’; ““>20 captures,’’ not “‘more than 20 captures’’). Use = 


for “‘equal to or less than’’ or “‘not more than” and = for “‘equal to or 
greater than”’ or “‘at least’? with numerals. 

When giving ranges, use “from 10.1 to 31.0 mm”’ or “ranges from 10.1— 
31.4,”’ but “‘the range is 21.3 mm.”’ In other words, do not use “‘to”’ unless 
it is preceded by “‘from.”’ 

When presenting equations and formulas, use the solidus (/) for simple 
fractions and give the meanings of all symbols and variables in the text. 
When presenting values with respect to another factor such as time or 
space, use the solidus (/) if only two measurements are involved (g/ha); if 
three or more measurements are involved use the exponent —1 as in “ 


g 


43 mg g'h"!.” Do not present equations published elsewhere unless they 
have been modified; a simple reference will do. 

In reporting measures of central tendency (means, modes, etc.) or 
dispersion (standard deviations, standard errors), units of measure should 
not be more precise than the original measurements. For example, if 
mammals are weighed to the nearest 0.1 g, then means such as 54.56 should 
be reduced to 54.6 and X + SE should be 54.6 + 0.1, not 54.6 + 0.09, 
both in tables and in text. 

Dental formulas should be presented as I 1/1, c 0/0, p 1/0, m 3/3, total 
18. Vertebral formulas should be presented as 7 C, 13 T, 6 L, 2-3 S, and 
26—31 Ca, total 55—60. Upper teeth should be referred to by capital letters 
and lower teeth by lowercase letters (e.g., P4 is the fourth upper premolar 
and ml is the first lower molar); do not use superscript and subscript 
numerals to designate upper and lower teeth. 

Time and dates.---Write dates as 24 April 1989, with no internal 
punctuation; an exception is in the specimen examined section where the 
three-letter month code should be used. Indicate time of day on the 24-h 
system with four digits. Midnight is written as 0000 h, 8:30 a.m. as 0830 
h, and 11:15 p.m. as 2315 h. Also, indicate time similarly as “4-h 
intervals,’ and “3 h/day.’ To express the age of a stratigraphic unit or the 
time of a particular geologic event, and where a specific dating technique 
has been used, employ “Ma” for mega-annum (time greater than one 
million years), or “B.P.” (years before present) for radiocarbon dated 
material, which goes up to 40,000 yrs. To express other time relations, use 
““m.y.”? (million years) or m.y.a. (million years ago). For example, from 50 
to 25 Ma, 25 m.y. elapsed and it is possible that these organisms lived at 
least 40 m.y.a. The ratio of light (L) hours to dark (D) hours under 
laboratory conditions is to be shown in this form: 14L:10D. 

Units of measure.---Use of the metric system and the international 
system of units (Systeme International d’Unités) is advocated. Exceptions 
to the use of the metric system and the international system of units are 
English units in localities from specimen labels and in quotations from 
other publications. Also, the following variables may be reported in other 
units: 


Temperature in degrees Celsius (C) instead of Kelvin (°K). 

Time in minutes (min) and hours (h) instead of seconds (sec). 

Sound intensity (relative terms) in decibels (dB) instead of 
watts/meter square (W/m7). 

Volume in liters (1) instead of cubic decimeters (dm‘°). 

Area in hectares (ha) instead of 10* m ?. 


Accepted abbreviations that can be used in text, tables, and figures of 
manuscripts, without explanation or punctuation, and some words that 
should be spelled out follow: 


Units of measure (use abbreviations only with numerical values, 


otherwise spell out; e.g., “... FN = 72-76, but populations with low 
fundamental numbers ... ’’): 

rE degrees Celsius 

2N (not 2n) diploid number (N = chromosome number) 

FN fundamental number 

n nautical 

g gravity 

Sec second 

min minute 

h or hr hour 

d day 

wk week 

mo month 

yr year 

m.y. million years 

g gram (not gm) 

mg milligram 

kg kilogram 

Hz hertz 

kHz kilohertz 

MHz megahertz 

km kilometer(s) 

m meter(s) 

mm millimeter(s) 

cm centimeter(s) 

fm fathom(s) 

nm nautical mile 

mi mile(s) 

ft foot (feet) 

in. inch (note period) 

diam diameter 

ha hectare 

M molar 

ppm parts per million 

W watt (or west) 


xX magnification 


Lg microgram 


pm micrometer (micron) 

gal gallon 

ml milliliter and cubic centimeter 

liter* should be spelled out when confusion can 


exist; mainly when it is used alone. When 
in print, “101” looks like a poorly spaced 
101. Confusion does not exist in a list of 
ingredients [(i.ec., 14 kg sawdust, 500 g 
potassium nitrate, 1.5 | water)] or when in 
combination (ml). 


Miscellaneous: Standard abbreviations used in text, their proper 
punctuation, and some terms that should be spelled out. 


AC DC alternating current and direct current 
B.A. B.S. M.A. 
M.S. M.Sc. Ph.D. college degrees 
Co; county 
counties spell out (no abbreviation) 
elev. elevation (note period) 
Fig. or fig. figure 
maximum no abbreviation 
minimum no abbreviation 


pers. comm. 
PO. 


personal (oral) communication 
post office 


R/V Searcher name of research vessel is underlined 
SEM scanning electron microscope 

States spell out to avoid confusion 

Table write out completely 

TEM transmission electron microscope 

UK United Kingdom (no spaces) 

U.S.A. United States of America 

USNM National Museum of Natural History 
weight no abbreviation 


Directions and coordinates: 


NES W ENE 
SE SSW (no periods) 
TI4N, R1OW, SW% 
Sec. Z legal description for localities 


10°06’N, 25°07’W latitude and longitude 


Latin and other foreign language terms (the following are accepted as 
written, without underlining; definitions follow terms). 
aff. affinis, having affinity with but not 
identical with 


a posteriori known from experience 


a priori intuitively, independent of experience 

ad lib. ad libitum, freely available 

auct. auctoris, of the author 

auctt. auctorum, or authors 

ca. circa, about 

cf. conferre, compare 

e:0%, (note comma) exempli gratia, for example 

emend. emended, emendation 

en masse in a body, as a whole 

et al. et alia, and others 

etc. et cetera, and so on 

neu (note comma) such as 

in litt. in a letter (Note: data or information in 
recent unpublished manuscripts should not 
be cited in any form; however, quotations 
On use ‘of data extracted frompyold 
manuscripts, as exist in archives, libraries, 
and occasionally herbaria, may be used) 

in situ in place 

in utero in the uterus 

in vitro outside the living organism 

in Vivo in the living organism 

leg. collector or collected by 

nec not 

nom. dub. nomen dubium, name of doubtful application 

nom. nud. nomen nudum (plural, nomina nuda), name 
without validation 

part. partim, part 

per se by itself, as such 

p-p. pro parte, in part 

sensu as defined by 

SIC thus (to signal exact transcription) 

Sell sensu lato, in the broad sense 

S.S. sensu stricto, in the strict sense 

via by way of, by means of 

viS-a-VI1S in relation to, as compared with 

VIZ. videlicet, namely 


vs. OF Versus 


against, in contrast to 


Statistical terms: 


ANOVA analysis of variance 

CV coefficient of variation 

df degrees of freedom 

n sample size or number in sample 
P probability 

rorR correlation coefficient 

SD standard deviation 

SE standard error 

SEM standard error of the mean 
¢ it UZ Statistical tests 

xX mean 


Symbols.---Male (d)) and female (2) symbols should not be used in the 

text, but may be used in figures and lists of specimens examined (8 ¢ or 
14 2 °—note space between number and symbol). Write percent as one 
word in the text, but use the percent sign after numerals (1%, 99%) and in 
bodies of tables. 
Because of possible confusion with similar symbols, do not use X as a 
symbol for the word “by.” Write, for example, ““Traps used were 7.6 by 
7.6 by 22.8 cm....” If a multiplication sign is used in mathematical 
formulae or for indications of magnification, identify it as such in the 
margin of the manuscript. 


Documentation and Literature Citation 


The function of literature citation is to assist readers in locating material - 
referenced by the author, a process that permits an orderly growth of 
knowledge through continued testing and reassessment. Documents written 
primarily to fill administrative requirements are not catalogued in most 
libraries and do not enter the body of knowledge that supports research. 
Therefore, such documents are not to be included in the Literature Cited 
section; the quarterly reports of U.S. Fish and Wildlife Service Cooperative 
Wildlife Research Units and job completion reports for Pittman-Robertson 
Federal Aid in Wildlife Restoration projects are examples of this kind of 
material. Certain other state, provincial, and federal reports also are 
excluded from lists of citations. Abstracts of oral presentations delivered 
at professional meetings and printed separately are excluded, but references 
-to abstracts in Dissertation Abstracts and Masters Abstracts and abstracts 
published in journals (e.g., American Zoologist, American Journal of 
Botany) are permitted. 


Style of documentation.---When citing informational references in text, 
use the form “Jones (1983)” if the author’s name is part of the sentence 
and “‘(Jones 1983)” if it is not. Two articles by one author cited at one 
time should be written “(Jones 1975, 1982)’’; two articles published by the 
same author in the same year, “(Jones 198la, 1981b).”’ Other examples 
follow: 

(Cameron 1977:507) or Smith (1957:23, table 3) [Note: only cite 
pagination in text references for direct quotations, in a synonymy, or in 
reference to a specific table or figure in another publication.] In text, 
citations of figures or tables other than those in the present manuscript 
should be in lowercase letters. Citations of figures or tables in the text 
should begin with a capitol letter, as: Fig. 1 or Table 2. 

‘In press”’ citations in text should not be used. Use the year of expected 
publication (current or subsequent year) and end the citation listing in the 
Literature Cited section with “‘(in press)’’—see Appendix II. This allows 
for less costly corrections when the citation is updated at the page proof 
stage. 

Citations with more than two authors are cited in the text as: Lidicker 
et al. (1976). Do not underline the “et al.’? Multiple citations in the text 
should be ordered chronologically and then alphabetically if in the same 
year, for example (Jones 1961, 1963; Hennings & Hoffmann 1977, Phillips 
1978, Jones & Smith 1981, Jones & Baker 1983, Mares et al. 1983) (Note: 
a semicolon should be used after citing two or more articles by the same 
author in a multiple citation, however, a comma should be used after 
different authors). Use an ampersand (&) between surnames of authors 
rather than “and.” In the text, multiple citations of three or more authors, 
in the same year with the same first author, should cite all authors, for 
example (Jones, Smith, & Williams 1988; Jones, Williams, & Smith 1988; 
Jones, Williams, Smith, & Baker 1988). 

In the text, unpublished material can be referenced as follows: (pers. 
comm.) denotes information obtained orally; (in litt.) denotes information 
obtained in a letter. Names of persons providing unpublished information | 
should include initials when referenced in text (e.g., R. H. Tamarin, pers. 
comm.; D. P. Christian, in litt.). In the Literature Cited section, do not cite 
or use information from unpublished recent manuscripts (except theses and 
dissertations) or papers in preparation. Unpublished data may not be 
referenced in any context. 

Citations in the text that indicate the author(s) of zoological scientific 
names should be as follows: Family Caryophyllidae Dana, 1846; Didelphys 
microtarsus Wagner, 1842:359; and, Themiste dyscrita (Fisher, 1952). Note 
the use of a comma after the author(s) name(s). In botanical scientific 


15 


names, dates may be used in the case of homonyms. The proper form for 
new zoological descriptions, plus their associated synonymy, is as follows: 


Chaceon bicolor, new species 
Figs. 1--3 


Geryon affinis.---Griffin & Brown, 1976:256, figs. 7--9.--- 
Sakai, 1978:9, figs. 18--19, pl. 2, fig. D (color). [Not 
Geryon affinis A. Milne Edwards & Bouvier, 1894] 


A more complete synonymy, with style examples, is as follows: 


Chrysogorgia spiculosa (Verrill, 1883) 
Figs. 5--6 


Dasygorgia spiculosa Verrill, 1883 (in part: 1 of 2 specimens 
from BL-190, 1 of 2 specimens from BL-195, not BL- 
205,? BL-222,? BL227), 23--24, pl. 2, fig. 5.---not 
Wright & Studer, 1889:9--10 (= C. affinis/elegans). 

Chrysogorgia sp. Agassiz, 1888:144, fig. 456.---Bayer, 1973: 
me, 1S. 

Chrysogorgia spiculosa: Versluys, 1902:53, 60, 61.---not 
Nutting, 1908:591.---Kiikenthal, 1919: 537, fig. 235; 
1924:406--407.---Bayer & Macintyre, 2001:342 (mi- 
nerology). 

Chrysogorgia elegans: Deichmann, 1936:231--232 (in part: 
pl. 22, fig. 7, pl. 33, fig. 1: BL-44 Gn part), BL-190, BL- 
195 Gn part), BL-200, and MCZ 4861). 


Note the names of subsequent users are separated from the name of the 
taxon by a distinctive and explicit manner (Article 51.2.1, International 
Commission on Zoological Nomenclature 1999), in this case by using a 
colon (:) or a period and a l-em dash (.---). 

The proper form for new botanical descriptions, plus their associated 
synonymy, is as follows: 


Pityrogramma mortonii Jackson, nom. nov. 


Pityrogramma mutabilis Gleas., Proc. Biol. Soc. Wash. 46: 
278. 1933, 

non L., 1753. Type: U.S.A.: Arizona: Cochise Co.: 7 mi NW 
of Wigwam, 4800 ft elev, Brooks 482 (US; isotype UC). 


All citations of author and date, whether informational, taxon describers, 
or essential references in the synonymy, except for botanical synonymies, 
must be listed in Literature Cited. Literature in botanical synonymies should 
be abbreviated according to the Botanico-Periodicum-Huntianum or 
Taxonomic Literature, ed. 2. 

The list of references at the end of the manuscript should have the 
primary heading “‘Literature Cited.’’ Sample literature citations are listed 
in Appendix II. Like all other parts of the manuscript, this section must be 
double-spaced. Only papers referred to in the text may be listed. The list 
must be alphabetical by authors’ last names. Therefore, within this 
structure, papers with one author would be listed first, then those with two 
authors (alphabetical by second author), then three authors, and so on. Use 
first author et al. (e.g., Patton, J. L., et al.) for papers having seven or more 
authors. Where the author line is identical, the listing should be 
chronological by publication date. If two or more papers by the same author 
or sequence of authors are listed, the name(s) are not repeated but are 
replaced by a 3 em dash (six dashes in typescript) and a period. Use a 3 
em dash in subsequent entries for all repeated authors. For example: 


Cockrum, E. L. 1974. 


------ Oc IK ones. inc 973: 
Leo »---===, & J. D. Smith: 1980. 


(Note: Throughout the text and in the Literature Cited, insert a space 
between the initials of a persons name, as above.) 

The volume number of a journal or other serial publication should be 
cited. However, do not include the issue number for journals having 
continuous pagination throughout the year or volume. When citing 
publications that have only a number and no volume, treat the number as ~ 
a volume (Occasional Papers of the Museum of Natural History, University 
of Kansas 25:1—39). Underline scientific names or other words only if 
italicized in the original title. In citing books, omit Roman pagination (e.g., 
“J. Wiley and Sons, New York, 432 pp.” not “*xxii + 432 pp.’’) except in 
instances that those pages contain the first reference to a taxon under 
consideration. Also, provide only the first city listed for publishers (e.g., 
John Wiley and Sons, New York’ not John Wiley and Sons, New York, 
Chichester, Brisbane, and Toronto’’). Plates (or figures), if not included in 
paginated materials, should be referenced after pagination (e.g., 286 pp. + 
pls. 1—24 [or figs. 1-24]). If plates, figures, or tables are paginated, do not 
cite them again in the reference. Journal names and book titles are set in 
Roman type, so do not underline. 


17 


Abbreviations used in Literature Cited.--- Abbreviations must not be 
used in the Literature Cited, except for those used in the title of the cited 
paper. Include and spell out all words that comprise the reference, including 
articles, conjunctions, and prepositions. Manuscripts will be returned to the 
author(s) if journal names are not completely spelled out. 


Conclusions 


Before submitting your manuscript, carefully check all citations in the 
text, including figure and table captions, against listings under Literature 
Cited, and check each entry in the Literature Cited section against its 
original source to verify title, year of publication, names, quotations, and 
page numbers. Include all citations, when appropriate, in synonymy 
listings. The editors scan the Literature Cited and make spot checks for 
accuracy, but cannot assume responsibility for verifying all citations. 

A summary in French, German, Russian, or Spanish is acceptable. This 
summary, which may be a translation of the abstract, should be placed 
immediately after the English abstract. Do not include an additional 
Summary Or Summary paragraphs in English at the end. 

The sequence of material in each copy should be: Title, Author(s), 
Author(s) Address(es), Abstract, Text, Acknowledgments, Literature Cited, 
Appendix, Figure Legends, Figure copies (each numbered and identified), 
Tables (each table numbered with an Arabic numeral and with heading 
provided). Number every page sent with the manuscript, including 
Literature Cited, figure legends, and tables. Any appendices should appear 
after Literature Cited. Figure legends and then copies of the figures follow. 
Next, the tables, each on a separate sheet. Glossy prints of each figure (or 
original artwork if high-contrast photographs of graphics cannot be 
obtained), should complete the parcel of manuscript materials. Three copies 
of all materials, including figures (legible photocopies acceptable), should 
be submitted with a cover letter stating the title, full name(s) of author(s), 
and availability of publication funds. Do not submit manuscripts under 
consideration for publication elsewhere. Author(s) address(es) must not 
have any abbreviations, including state or province names. Also, indicate 
country, including U.S.A., when appropriate. 


Acknowledgments 


The acknowledgments section should contain a few brief statements in 
a single paragraph to recognize the contribution of others and support from 
patrons or agencies. Use only initials for names of persons acknowledged, 
but spell out names of agencies (e.g., National Science Foundation not 
NSF). Parts of this manuscript were extracted intact and other parts 


18 


paraphrased from a similar manuscript prepared by D. E. Wilson, A. L. 
Gardner, and B. J. Verts for The Journal of Mammalogy, from “‘Standard 
style for technical publications’? prepared for customers of Allen Press, 
Inc., by G. Dresser, and “‘The Allen Press Guide to Illustration 
Preparation.’ Helpful suggestions for this publication were provided by C. 
C. Baldwin, S. D. Cairns, E D. Ferrari, S. L. Gardiner, G. R. Graves, G. 
D. Johnson, R. Lemaitre, R. B. Manning, W. N. Mathis, and D. E. Wilson. 


Literature Cited 


Costain, D. C. 1978. Dynamics of a population of Belding’s ground 
squirrels in Oregon. Unpublished M.S. Thesis, Oregon State 
University, Corvalis, 66 pp. 

Couch, L. K. 1932. Breeding notes on a few Washington mammals. ---The 
Murrelet 13:25. 

Day, R. A. 1994. How to write & publish a scientific paper. Fourth edition. 
Oryx Press, Phoenix, Arizona, 223 pp. 

International Commission on Zoological Nomenclature. 1999. International 
code of zoological nomenclature, fourth edition. The International 
Trust for Zoological Nomenclature, The Natural History Museum, 
London, 306 pp. 

Junge, R., & D. E Hoffmeister. 1980. Age determination in raccoons from 
cranial suture obliteration.---The Journal of Wildlife Management 
44:725-729. 

Linsdale, J. M. 1946. The California ground squirrel: a record of 
observations made on the Hastings Natural History Reservation. The 
University of California Press, Berkeley, 475 pp. 

McKeever, S. 1966. Reproduction in Citellus beldingi and Citellus lateralis 
in northeastern California. Pp. 365-385 in I. W. Rolands, ed., 
Symposia of the Zoological Society of London, Academic Press, 
London 15:1—559. 

Scheffer, T. H. 1941. Ground squirrel studies in the Four-Rivers country, 
Washington.---Journal of Mammalogy 22:270—279. 

Style Manual Committee. 1994. Scientific style and format: the CBE 
manual for authors, editors, and publishers. Sixth edition, Council 
of Biological Editors, Inc., Cambridge University Press, N.Y., 825 


Pp. 
Tomich, P. Q. 1962. The annual cycle of the California ground squirrel. 
---University of California Publications in Zoology 65:213—281. 
Wilson, D. E., A. L. Gardner, & B. J. Verts. 1989. Guidelines for 
manuscripts for publications of the American society of 
mammalogists.---Journal of Mammalogy (supplement) 70(4):1—17 
+ 21 unnumbered. 


Appendix I 


Examples of Typescript for Page Having Figure Legends 


Fig. 1. Localities at which Sorex trowbridgii (open circles) and S. 
bendirii (closed circles) were collected in Benton Co., Oregon, 1975-1986. 


Fig. 2. Relationship between body mass and body length for (a) 12 Sorex 
trowbridgii, and (b) five S. bendirii collected in Benton Co., Oregon, 1975— 
1986. 


Fig. 3. Syringonomus dactylatus. A. Photomicrograph of the cervical 
region of the holotype, USNM 77172; unlabeled arrows point to amphidial 
gland; scale equals 100 wm. B. Photomicrograph of the posterior body 
region and tail of the paratype USNM 77175; scale equals 100 pm. 
Abbreviations.---AV, anal vent; CG, caudal gland; NR, nerve ring. 


Appendix II 


Examples of Literature Citations, with Parenthetical Comments on Style 
Following Some Entries 


Journals. --- 


Bader, R. S. 1965. A partition of variance in dental traits of the house 
mouse. ---Journal of Mammalogy 46:384—388. 

Blair-West, J. R., et al. 1968. Physiological, morphological and behavioural 
adaptation to a deficient environment by wild native Australian and 
introduced species of animals.---Nature 217:922—928. [use of “‘et 
al.” for papers with seven or more authors] 

Blake, J. A. 1985. Polychaeta from the vicinity of deep-sea geothermal 
vents in the eastern Pacific I: Euphrosinidae, Phyllodocidae, 
Hesionidae, Nereididae, Glyceridae, Dorvilleidae, Orbiniidae, and 
Maldanidae. Pp. 67—101 in M. L. Jones, ed., Hydrothermal vents of 
the eastern Pacific: an overview.---Bulletin of the Biological 
Society of Washington 6:1—566. 

Carleton, M. D., & C. B. Robbins. 1985. On the status and affinities of 
Hybomys planifrons (Miller, 1900) (Rodentia: Muridae). --- 
Proceedings of the Biological Society of Washington 98:956—1005. 
[Underlined scientific names will be set in italics; note use of an 
ampersand (&) instead of “‘and’’ between authors’ names] 

Fauvel, P. 1936. Sur quelques annélides polychétes de I’Ile de Pacues.--- 
Bulletin du Muséum National d’ Histoire Naturelle, Série 2, 8:257— 
259. [Include all diacritical marks] 


20 


Ortiz, D. L., P. Costa, & B. J. LeBoeuf. 1990. Water and energy flux in 
elephant seal pups fasting under natural conditions. --- Physiological 
Zoology (in press). [Note use of probable year of publication after 
authors names and the use of “in press,” in parentheses, after the 
journal name] 

Whitaker, J. O., Jr, & R. E. Wrigly. 1972. Napaeozapus insignis. --- 
Mammalian Species 14:1—6. [underline scientific names only if 
italicized in original title] 

Wicksten, M. K. 1984. Alpheopsis harperi (Decapoda: Alpheidae): A new 
species of snapping shrimp from Texas.---Northeast Gulf Science 
7(1):97—100. 


Proceedings and transactions. --- 

Chitty, D. 1952. Mortality among voles (Microtus agrestis) at Lake 
Vyrnwy, Montgomeryshire in 1936—9.---Philosophical Transactions of 
the Royal Society of London, Series B, 236:505—5572. 

Kirsch, L. M., & A. D. Druse. 1973. Prairie fires and wildlife.--- 
Proceedings of the Annual Tall Timbers Fire Ecology Conference 12: 
289-304. 

Peek, J. M., M. C. Scott, L. J. Nelson, D. J. Pierce, & L. L. Irwin. 1982. 
Role of cover in habitat management for big game in northwestern 
United States.---Transactions of the North American Wildlife and 
Natural Resources Conference 47:363-—373. 


Books. --- 


Baird, S. F 1857 [1858]. Mammals. in Reports of explorations and surveys 
for a railroad route from the Mississippi River to the Pacific Ocean. 
Beverly Tucker, Printer, Washington, D. C., 8(part 1):1—757 + pls. 17— 
28, 30—60. [give correct publication date in brackets] 

Barbour, R. W., & W. H. Davis. 1969. Bats of America. The University of 
Kentucky, Lexington, 286 pp. [City is not followed by state when 
name of state is part of the name of the press] 

Eisenberg, J. EK 1981. The mammalian radiations: an analysis of trends in 
evolution, adaptation, and behavior. The University of Chicago Press, 
Chicago, 610 pp. [Names of large, universally recognized cities are 
not followed by state] 

Honacki, J. H., K. E. Kinman, & J. W. Koeppl (eds.). 1982. Mammal 
species of the world: a taxonomic and geographic reference. Allen 
Press, Inc. and The Association of Systematics Collections, Lawrence, 
Kansas, 694 pp. 

Hsu, T. C., & K. Benirschke. 1969. Microtus oregoni (creeping vole) 2n = 


21 


17, 18. An atlas of mammalian chromosomes. Vol. 3, Folio 121, 
Springer-Verlag, New York, unpaged. [Pages are not numbered] 
Linnaeus, C. 1758. Systema nature per regna tria naturae, secundum 
classes, ordines, genera, species, cum characteribus, differenti, 
synonymis, locis. Tenth edition. Laurentius Salvius, Stockholm 1:1-— 
824. [give complete title, do not use ellipsis] 

Nowak, R. M., & J. L. Paradiso. 1983. Walker’s mammals of the world. 
Fourth edition. Johns Hopkins University Press, Baltimore, 1 and 2: 
1—1362. 


Part of book.--- 


Bourliere, EF 1955. Ordre des Fissipedes. Systématique. Pp. 215-291 in P- 
P. Grasse, ed., Traité de Zoologie. Masson et Cie., Paris 17(1):1—1170. 
[period after title of volume] 

Calder, W. A., II. 1974. Consequences of body size for avian energetics. 
Pp. 86—151 in R. A. Paynter, Jr, ed., Avian energetics. Publication of 
the Nutall Ornithological Club, Cambridge, Massachusetts 15:1—334. 

Carleton, M. D., & G. G. Musser. 1984. Muroid rodents. Pp. 289-379 in 
S. Anderson & J. K. Jones, Jr., eds., Orders and families of Recent 
mammals of the world. John Wiley and Sons, New York, 686 pp. [give 
pagination for entire volume] 

Leach, W. E. 1821. Galatéadées. Pp. 49-56 in E G. Levrault, ed., 
Dictionnaire des Sciences Naturelles, 18(1820), Strasbourg. 

Spezharsky. T. N. 1939. Order Ostracoda. Pp. 193-196, pl. 46 in B. 

-Likharev, ed., Atlas rokovodyashchikh form iskopaemykh faun SSSR. 
Volume 6, Permskaya Systema. Tsentral’nyi Nauchno-Issledova- 
tel’skiy Geologo-Razvedochnyi Institut (TsNIGRI). (The atlas of the 
leading forms of the fossil fauna USSR. Volume 6, Permian System. 
Central Geological and Prospecting Institute) (¢n Russian). 


Theses and dissertations. --- 


Lackey, J. A. 1973. Reproduction, growth, and development in high- 
latitude and low-altitude populations of Peromyscus leucopus 
(Rodentia). Unpublished Ph.D. dissertation, University of Michigan, 
Ann Arbor, 128 pp. 

Wrazen, J. 1976. Feeding ecology of a population of eastern chipmunks 
(Tamias striatus) in southeast Ohio. Unpublished M.S. thesis, Ohio 
University, Athens, 26 pp. 


Table 1.---Some reproductive patterns in four species of Spermophilus in western 
United States. 


Litter size? 


Species and pola | Was nA Shee oe Reproductive 
state n x Range season? Authority 

S. washingtoni 

Washington 26 8.0 5-11 February—March Scheffer 1941 
S. beldingi 

Oregon 110 3 5+8 April—June Costain 1978 

California 37/ Vail 3-10 April—July McKeever 1966 
S. columbianus 

Washington 21 5.8 2-7 March—May Couch 1932 
S. beecheyi : 

California 40 TD 5-12 February—April Linsdale 1946 

California 34 6.1 1-9 March—June Tomich 1962 


4 Based on counts of embryos. 
> From first evidence of reproduction behavior to first appearance of young. 


Table 2.---Means (+=SE) and ranges for mass and length of 514 bacula from raccoons 
(Procyon lotor) among five age classes, northwestern Oregon, 1982—1983 and 1983-1984 
furbearer seasons. 


Age Baculum mass (g) Baculum length (mm) 
class? n 2S SE SIE, Range X + SE Range 
1 205 0.63 + 0.01 0.30—1.22 (ESS 05 54.5-88.9 
2, 145 2.99 + 0.04 2.03-4.16 105.9 + 0.4 92.7-117.5 
3 48 3169) ==. 0108 2.63-4.76 HOWE) Se OLS) 102.6—117.7 
4 43 4.01 + 0.07 2.76-4.86 109.3 = 0.6 102.1—119.5 
a) 73 4.40 + 0.07 3.05—5.67 NOY 2 28.055 99.0-118.8 


@ Age classification based on cranial-suture technique (Junge & Hoffmeister 1980). | 
> Includes all individuals =4.5 years old. 


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